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Malesian Euphorbiaceae Descriptions |
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66. MACARANGA (Euphorbiaceae)
T.C. Whitmore (primary author) plus editors M.J.E. Coode & Petra Hoffmann
S.J. Davies (sections Pachystemon, Pruinosae & Winklerianae)
J.W.F. Slik, Priyono & P.C. van Welzen (Borneo species)
W. Takeuchi (Macaranga daviesii)
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Whitmore's book.
Davies, S.J. 2001. Systematics of Macaranga sect. Pachystemon and Pruinosae (Euphorbiaceae). Harvard Pap. Bot. 6: 371448.
Slik, J.W.F., Priyono & P.C. van Welzen. 2000. Key to the Macaranga and Mallotus species (Euphorbiaceae) of East Kalimantan, Indonesia. Gard. Bull. Singapore 52: 1187.
Takeuchi, W. 2007. Additions to the Flora of the Kaijende Highlands, Papua New Guinea: Macaranga daviesii (Euphorbiaceae), a new calciphilous species from the Paiela limestone. Harvard Pap. Bot. 12: 389394.
Whitmore, T.C. 2008. The genus Macaranga, a Prodromus. Royal Botanic Gardens Kew.
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Infrageneric and Informal groups:
Dioica group (plus New Guinean Pseudorottlera)
Regional Keys to the species
Key to the species the Malay Peninsula
Key to the species of Java and Bali
Key to the species of the Philippines
Key to the species of Sulawesi (Celebes)
Key to the species of the Lesser Sunda Islands (minus Bali) and the Moluccas
Key to the species groups in New Guinea
Inadequately known Macaranga names
Macaranga Thouars, Gen. Nov. Madagasc. (1806) 26; Mόll.Arg. in DC., Prodr.15, 2 (1866) 987; Prain, Fl. Trop. Afr. 6(1) (1912) 932 Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 298; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 434; Backer & Bakh.f., Fl. Java 1 (1963) 487; Airy Shaw, Kew Bull. 26 (1972) 286; Whitmore, Tree Fl. Malaya 2 (1973) 105; Airy Shaw, Hooker's Icon. Pl. 38 (1872) t. 3718; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975), 140; Coode, Taxon 25 (1976) 184; Whitmore in Airy Shaw, Kew Bull. Addit. Ser. 8 (1980) 123; Airy Shaw, Kew Bull. 35 (1980) 646; Whitmore, Kew Bull. 36 (1981) 312; Airy Shaw, Kew Bull. 37 (1982) 26; Alph. Enum. Philipp. Is. (1983) 33; G.L.Webster, Ann. Missouri Bot. Gard. 81 (1994) 86; Govaerts, Frodin & Radcl.-Sm., World Checkl. Bibliogr. Euphorb. 3 (2000); Radcl.-Sm., Gen. Euphorbiacearum (2001) 213; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 361; Whitmore, Gen. Macaranga Prodr. (2008) 1; G.L.Webster in Kubitzki, Fam. Gen. Vasc. Pl. 11 (2014) 127. Lectotype species (designated by Coode 1976): Macaranga mauritiana Bojer ex Mόll.Arg.
Panopia Thouars, Gen. Nov. Madagasc. (1806) 26, nomen tantum.
Mappa A.Juss., Euphorb. Gen. (1824) 44; Baill., Ιtude Euphorb. (1858) 428. Type species: Mappa glabra A.Juss. [= Macaranga glabra (A.Juss.) Pax & K.Hoffm.].
Pachystemon Blume, Bijdr. (1826) 626. Type species: Ricinus trilobus Reinw. ex Blume = Pachystemon trilobum (Reinw. ex Blume) Blume [= Macaranga triloba (Reinw. ex Blume) Mόll.Arg.].
Mecostylis Kurz ex Teijsm. & Binn., Natuurk. Tijdschr. Ned.-Indiλ 27 (1864) 27. Type species: Mecostylus acalyphoides Kurz ex Teijsm. & Binn. [= Macaranga involucrata (Roxb.) Baill.].
Phocea Seem., J. Bot. 8 (1870) 69. Type species: Phocea andersonii Seem. [ Macanga sp.].
Trees, usually small to 15 m, occasionally to 20(30) m tall, or treelets as small as 2 m, rarely shrubs or lianas, sometimes sarmentose; dioecious; trunks and twigs sometimes armed with spines; crown usually becoming sympodial, with few orders of branches. Bark sometimes prominently hooped (rings surrounding the stem); wood soft. Indumentum, if present, of simple or rarely of tufted hairs. Twigs slender to stout, in a few species hollow and ant-inhabited. Stipules paired, or in a few species united and apparently single, entire, subulate to large and leafy, erect, spreading or recurved, often caducous, occasionally leaving conspicuous scars, with starchy granules (Beccarian bodies') in ant-species. Leaves spirally arranged, simple, or sometimes 3(7) lobed; petioles short to longer than leaf, sometimes kneed at one end or both; blades papery to leathery, base cuneate, rounded, cordate or cordulate (the lobes sometimes overlapping) sometimes caudate, sometimes peltate, sometimes with glands on adaxial surface on main nerves or lamina or margin at the base near petiole insertion; margin sometimes with protruding variously conspicuous glandular teeth often terminating secondary nerves, apex usually acute or acuminate, adaxial surface usually glabrous, occasionally hairy on main nerves near base, abaxial surface nearly always with tiny golden to black usually dense or sometimes sparse granular glands, and glabrous to variously hairy on nerves or nerves plus lamina, rarely glaucous; usually pinnately nerved or either pinnately nerved but with midrib plus 2 main nerves at petiole insertion, or palmately nerved, with spider's web nervation and multiple nerves from petiole insertion. Inflorescences shorter to somewhat longer than leaves, axillary, amongst or behind leaves, spikes, racemes or panicles, with up to 4 axis orders, staminates usually more branched; first branches usually alternate or at least lowermost sometimes opposite, occasionally with accessory axillary branches; axes stout to slender to threadlike, occasionally with (sub)persistent usually small bracts subtending branches. Flowers in clusters of few to many, subtended by often persistent bracteoles, either shorter or longer than cluster, these clusters on last 1 or 2 axis orders, contiguous to distant and usually spiral; bracteoles minute to >1 cm and leafy, margin entire to dissected to laciniate and sometimes glanded, papery to fleshy, adaxially with 1-several small to large patellar glands; flowers minute, seldom >1 mm, sessile or pedicellate; sepals fused or 2 or 3; petals and disc absent; in staminate flowers stamens 1-many, anthers 24-locular (3- or 4-locular in Malesia), opening by splits, pistillode absent; in pistillate flowers ovary single, with 15 locules, each with 1 ovule, style(s) short or long, usually terminal, occasionally lateral or basal to ovary, stigmas often plumose. Fruits 15-locular, stalked or sessile; calyx persistent or not; wall leathery, fleshy or thinly to thickly woody, often waxy with granular glands, or glabrous, or occasionally hairy, smooth or with short to long close to distant slender to stout spines, or horned, sometimes with persistent style(s), ultimately dehiscent. Seeds: testa smooth to pitted to verrucose, arillate or not, aril red to pink.
Distribution A species rich genus of c. 280 species. About 50 species in Africa, Madagascar, Comoro Islands, and Mauritius. Most species, more than 200, are found in the tropical regions of Asia (India and Sri Lanka to southern China, Japan, Indochina, Taiwan, throughout Malesia) to the West Pacific islands and Australia. In Malesia c. 182 species.
Note There are still new species being discovered, especially in the rich regions of Borneo and New Guinea, even though these are well-collected. In central Malesia (Sulawesi, Moluccas, Lesser Sunda Islands), new species and range extensions are being discovered as these poorly collected regions become better collected.
Macaranga acerifolia Airy Shaw, Kew Bull. 33 (1978) 67; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 139, Pl. 6, fig. 1; Whitmore, Gen. Macaranga Prodr. (2008) 67. Type: BW (Schram) 15079 (holo K; iso L), Indonesia, Papua, Meos Num, Geelvink Bay.
Small tree. Twigs slender, c. 3 mm diam., angled near tip, finely furfurescent. Stipules narrowly elliptic, c. 3 by 0.8 mm, often curved, acute, finely pubescent, erect, a few persistent. Leaves: petioles 24 cm long, slender, terete, finely crisped-pubescent, not kneed; blades mostly broadly trilobed to 18 by 12 cm (a few elliptic, up to 14 by 2cm), papery, drying grey-green, lobes long-caudate, central lobe to 13 by 5 cm, sinuses very broad, lateral lobes divergent, narrow, to 7 cm long, 2cm broad at base, base broadly rounded, minutely cordate to peltate, with 13 glands on upper surface, margin entire, lower surface rusty furfurescent on lower part of main nerves, secondaries slender arching and joining near margin, basal pair extending into lateral lobes. Staminate inflorescences and flowers unknown. Infructescences slender, up to 12 cm long spikes with an apical cluster of leafy bracts, axis slender, terete, finely crisped pubescent; bracts ovate, to 15 by 7 mm, papery, granular glandular, margins coarsely acute to acuminate with large-dentate teeth with long hairs, apex acuminate, with midrib and secondary nerves with close short crisp hairs. Fruits globose, c. 6 by 5 mm, granular glandular and with sparse c. 1 mm long caducous spines, thinly woody; pedicel 13 mm long, densely puberulous; calyx persistent, < 1 mm high; style persistent, subapically recurved, densely plumose. Seeds spherical, coarsely verrucose, with sarcotesta.
Distribution Indonesian New Guinea (Geelvink Bay).
Habitat & Ecology Common in primary forest at sea level.
Macaranga advena Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv (1919) 31; L.M.Perry, J. Arnold Arbor. 34 (1953) 229; Airy Shaw, Kew Bull. 23 (1969) 110, passim; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 161; Whitmore, Gen. Macaranga Prodr. (2008) 67. Type: Ledermann 12490 (B?, lost).
Small tree 56 m. Twigs slender, ferruginous tomentose, becoming glabrous. Stipules ovate, c. 8 mm long, apex acuminate, glabrous, caducous. Leaves: petioles 23 cm long, slender, pilose; blades elliptic, 920 by 1.54 cm, leathery, base rounded, very narrowly cordate with the lobes overlapping, margin entire, apex caudate, extremely long and narrowly linear, c. 4 cm long, upper surface glabrous above, with several glands near base, lower surface sparsely pilose, densely granular-glandular; nerves 68 pairs, veins and veinlets reticulate. Staminate inflorescences and flowers unknown. Pistillate inflorescences racemes (? spikes), 510 cm long, axis slender tomentellose, bearing sterile bracts below; bracteoles rhombic ovate, c. 6 mm long, narrowed at base, marginal patelliform glands within, pilose and densely granular-glandular without. Pistillate flowers 3 or 4 crowded at tip, with calyx cupular, denticulate, granular-glandular, ovary 1-locular softly spiny, style lateral, 23cm long, densely plumose. Fruits and seeds unknown.
Distribution Papua New Guinea (Felsspitze).
Habitat & Ecology Forest clearings. Altitude: 14001500m.
Notes 1. Only known from the type collection, which was not seen by me. The description is paraphrased from Perry's translation, amended against the original Latin. The other species of this group have pistillate racemes, here the description is of spikes.
2. Airy Shaw draws a similarity to his M. misimae, with which I concur. That has similar staminate bracteoles, but differs conspicuously in its persistently pubescent twigs, and leaves 12-nerved and not caudate.
Macaranga aenigmatica Whitmore, Gen. Macaranga Prodr. (2008) 68 Type: Whitmore 3079 (holo K; iso BO, BZF), Indonesia, Sulawesi, South, Malili.
Small tree to 15 m. Twigs solid, stout to 10(15) mm diam., terete, drying coarsely shallowly striate, intensely glaucous, glabrous. Stipules long-deltoid, up to 30 by 15 mm, tapering from base to acute apex, somewhat hooded, dark brown but somewhat glaucous, glabrous, thinly leathery, recurved, long persistent. Leaves: petiole 2036 cm long, c. 5 mm diam., rather slender, terete, slightly striate, somewhat glaucous, glabrous; blades broad, trilobed (28)34(46) by (22)28 or more cm, thinly leathery, sometimes drying grey green above, base rounded to broadest point above petiole insertion, strongly (5)8(15)cm peltate, margin with conspicuous 1cm spaced protruding glands and often coarsely sinuous, central lobe c. 14 cm wide, lateral lobes c. 6 cm wide, wide sinuses c. 10 cm deep, apices acute to acuminate, lower surface glabrous or with sparse minute stellate hairs, with sometimes scattered, often inconspicuous dark granular glands, tertiary nerves conspicuous, raised below, strongly scalariform, quaternary nerves faint, scalariform. Staminate inflorescences axillary open panicles, up to 20 by 10 cm, 4 orders of opposite divaricate branches, axes angled and striate, glaucous, at first weakly rufous furfuraceous, basal peduncle 17 cm long, secondary branches c. 6 cm distant, quaternary c. 1 cm distant; bracts persistent, ovate, c. 2 by 1 mm, apex acute, clusters of 4 or 5 flowers grouped on ultimate branches; bracteoles bigger than cluster, persistent, oblong, c. 4 by 2 mm, apex acuminate, narrowed to base, central part concave, margin entire, glandless, finely pubescent on both surfaces. Staminate flowers c. 0.5 mm diam.; pedicel c. 0.5 mm long; stamens 1 or 2, anthers 4-locular. Pistillate inflorescences as staminate but only 3 branching orders. Pistillate flowers solitary or paired, calyx furfuraceous, ovary 2- or 3-locular, developing a stout furfuraceous pedicel in fruit. Fruits up to 1 by 1 cm, bi- or tricarpellate, loculicidally dehiscing, with a raised ridge at the sutures and grooved to lobed at the septa; wall woody, smooth, blackish brown, closely granular glandular, tipped by 3 caducous spreading, c. 1 mm long, smooth stigmas; pedicel stout, glaucous, 1520 mm long; calyx persistent. Seeds round, verrucose, with sarcotesta.
Distribution Endemic to Sulawesi.
Habitat & Ecology Lowlands, locally common. Altitude: up to 1500 m.
Note Highly distinctive from its twigs, stipules and fruits.
Macaranga aλtheadenia Airy Shaw, Kew Bull. 29 (1974) 322; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 145; S.J.Davies, Harvard Pap. Bot. 6 (2001) 384. Type: Chai S 19202 (holotype: K; isotypes: A, L, SAR, SING), Malaysia, Sarawak, Anap, Ulu Mayeng.
Macaranga calcifuga auct. non. (Whitmore) R.I.Milne, Kew Bull. 49 (1994) 453, p.p.
Tree to 15(25) m tall, 15(30) cm dbh; twigs 710 mm in diam., glabrous, densely glaucous, hollow, housing ants. Stipules ovate, 810 by 813 mm, recurved, succulent, producing food-bodies on the abaxial surface, 13 (rarely more) pairs present on the shoots. Leaves: petioles terete, slender, 1530 cm long, glaucous, glabrous; blades broadly to very broadly ovate, large, variable in size, 1540 by 1335 cm, usually shallowly trilobed, rarely with just three cusps or with an additional pair of tiny basal leaf cusps, dissected to Ό½ of the leaf length, central lobe c. 712 cm wide with an acuminate apex, lateral lobes ascending with acuminate apices, 25 cm peltate, base broadly to very broadly rounded, margin entire, with huge (23 mm long), prominent conical nectaries, particularly dense along the leaf base; adaxial surface glabrous, abaxial surface evenly covered in minute crisped silvery hairs and scattered with erect soft hairs, densely glaucous; venation palmate with 7(8) prominent 1° veins. 2° venation prominent and densely scalariform, looping and terminating at the margin in conical nectaries, 3°4° venation prominently scalariform; young leaves red-brown, densely covered in minute ferrugineus hairs. Staminate inflorescences particulate, erect, 1424 by 915 cm, mostly glabrous. distal axes finely ferrugineus pubescent, to 4 axis orders, main unbranched axis flattened and 36 cm long, first pair of secondary branches ±opposite with accessory branches; bracts elliptic, 58 by c. 5 mm, margin entire, apex acute, glabrous, caducous; flower clusters with c. 1520 flowers, decussately arranged and strongly grouped at distal ends of inflorescence branches; bracteoles ovate to broadly ovate, c. 4 by 45 mm, enclosing flower clusters, margin finely and evenly pectinate with c. 20 fine teeth each with minute erect hairs at their apices, apex rounded to acute, both surfaces glabrous, adaxial surface with a dense patch of minute red-brown hairs at base. Staminate flowers 101.5 mm long, shortly pedicellate, pedicel not persisting; sepals fused, splitting to c. 1/2, apex glabrous; stamens 13; anthers 3-locular. Pistillate inflorescences paniculate, erect, 714 by 410 cm, stout, glabrous or with ferrugineus hairs near apex, to 3 axis orders. secondary branches ± opposite; bracts elliptic, margin entire or finely pectinate, apex acute, mostly glabrous, caducous. Pistillate flowers c. 3.5 by 2 mm, solitary in bracteole axils; calyx urceolate, apex truncate, c. 2 mm long, glabrous, persistent, splitting irregularly: ovary 45-carpellate, c. 2 mm long; styles c.1.5 mm long, fused to c. 1/2 of the length, persisting to form a prominent c. 2 mm long crown at fruit apex; stigma not dissected. Fruits 57 by 811 mm, subglobose, ± sessile, one discrete glandular patch on each carpel wall developing into a slightly raised swelling, appearing almost flat when dry, covered in sticky exudate. Seeds subtriangular-ovoid. 3.54.0 mm in diam., black, with shallow well-spaced irregular grooves, encased in a fleshy bright red-pink aril.
Distribution Borneo (Brunei, East and West Kalimantan, Sarawak).
Habitat & Ecology A gregarious tree on relatively nutrient-rich soils in lowland forest to 900 m altitude. The species is commonly found on shale-derived soils in Kuala Belalong, Brunei, on basalt derived soils of Bukit Mersing in central Sarawak, and on shale-derived soils of the Setap Formation in northern Sarawak, such as in Lambir Hills and Gunung Mulu National Parks. It is a small- to medium-sized early successional tree occurring along river banks and in forest openings within primary forest. It also occurs on forest edges and along roadsides.
Notes 1. The enormous "volcanic" nectaries on the leaf margins are diagnostic for this species. However, these nectaries do not develop fully until the tree is more than 12 m tall.
2. Although this species comes out in the bancana-clade in the cladistic analysis, M. aλtheadenia shares a series of morphological characters with the species of the motleyana-clade. The staminate flowers are shortly pedicellate and have 13 stamens, the staminate bracteoles have pectinately dissected margins and have a decussate arrangement, and the abaxial leaf surfaces and twigs are strongly glaucous. These character states are also found in the motleyana-clade and suggest a closer relationship than is shown with the current cladistic analysis.
Macaranga albescens L.M.Perry, J. Arnold Arbor. 34 (1953) 244; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 139; Whitmore, Gen. Macaranga Prodr. (2008) 69. Type: Clemens 7550B (holo A), Papua New Guinea, Morobe Prov., Sarawaket.
Tree 1030 m high. Twigs stout, 34 mm diam., strongly angular, coarsely striate, pubescent. Stipules linear-elliptic, c. 4 mm long, pubescent, very soon caducous. Leaves: petioles 39 cm long, slender, 12 mm diam., terete, pubescent, kneed at both ends; blades ovate, 613 by 410 cm, leathery, drying grey brown above, pale brown often with whitish patches below, base 510 mm peltate, with a usually prominent protruding basal lobe to 6 mm wide, margin usually recurved, apex broadly acute, at first brown floccose above becoming glabrous, below evanescent brown floccose over a dense finely felted white tomentum and black granular glands then becoming glabrous almost glandless and shagreened; venation often coarsely to finely bullate, with 3 major and several minor nerves at petiole insertion, latter penetrating lobe, secondaries and reticulations slightly raised above. Staminate inflorescences sparsely branched racemes to 7 cm long, pubescent with a few granular glands, branches to 3 cm long, alternate, divaricate or ascending; peduncle 1015 mm long, main axis c. 1 mm diam., terete, striate, flower clusters in groups; bracteoles variable, oblong, blunt, c. 2 by 1 mm, or triangular, c. 1 by 1mm, margin irregularly coarsely dentate, papery, pubescent without, persistent, spreading, smaller than cluster. Staminate flowers c. 15 per cluster, shortly pedicellate, calyx pubescent and black granular glandular, stamens 46, anthers 4-locular. Infructescences open sparsely branched pubescent racemes to 13 cm long; peduncle 13 cm long, main axis c. 2 mm diam., angled, coarsely ridged. Fruits solitary or in small groups, ellipsoid, c. 4 mm long, leathery, densely pubescent and black granular glandular; pedicel 210 mm long, pubescent; calyx persistent; style single, c. 3 mm, lateral, curved, persistent, slightly papillose. Seeds globose, rugose, with sarcotesta.
Distribution New Guinea.
Habitat & Ecology Primary and disturbed montane forests. Altitude: 21002800 m.
Note Stipules described from Perry (1953). Staminate bracteoles variable. Leaves similar to M. trichanthera except for the basal lobe.
Macaranga aleuritoides F.Muell., Descr. Notes Papuan Pl. 2 (1876) 21; Pax & K.Hoffm. (1914) 366, t. 61 D,E; L.M.Perry, J. Arnold Arbor. 34 (1953) 204; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)139; Airy Shaw, Kew Bull. 37 (1982) 26; Whitmore, Gen. Macaranga Prodr. (2008) 71. Type: S. Macfarlane s.n. (holo MEL?), Papua New Guinea, Western Prov., Fly River.
Macaranga riparia Engl., Bot. Jahrb. 7 (1886) 463; K.Schum. & Lauterb., Fl. Schutzgeb. Sόdsee (1900) 397; J.J.Sm., Nova Guinea Bot. 8 (1910, 1912) 235, 787.
Tree to 24 m, often less. Twigs (4)10(16) mm diam., somewhat angular at least near tip, with conspicuous stipule scars, sometimes ridged, finely furfuraceous or glabrous. Stipules united, apparently single, enclosing a conspicuous erect, tapering terminal bud, 711 cm long, 0.8 by 3 cm wide at base, ovate-elliptic, acuminate, leathery but not rigid, finely golden tomentose usually densely so, subpersistent. Leaves: Petioles 1432 cm, usually stout, usually strongly D-shaped in transverse section near base and 411 mm across, at first tomentose, soon glabrous, drying constricted at base; blades large, strongly trilobed (rarely shallowly lobed or entire ovate), 2232 by 2027 cm, sinuses broad, penetrating blade to 1/3 or less, all lobes acute, all broad (the central most so), central lobe largest, base rarely truncate usually (deeply) cordate with a (deep) narrow square-bottomed central sinus, at petiole insertion 2 rounded auricles with on upper surface several large glands, drying khaki brown, margin repand with distant small slightly protruding glandular teeth, leathery, above glabrous shiny, below all nerves slightly raised and finely pubescent black granular glandular, 3 main and several minor nerves from petiole insertion, tertiaries and quaternaries reticulate. Staminate inflorescences dense, crowded, 20 by 8 cm, dirty-brown pubescent, slender branched panicles with 3 axis orders, main axis flattened near base to 5 mm wide, strongly tapering upwards, peduncle to 2.5 cm, most branches opposite, a few with axillary branches, ultimate branches sinuous, thread-like; bracts ovate oblong, 15 by 5 mm, apex acute tomentose, soon caducous; bracteoles linear, 2 mm, apex acuminate, slightly swollen with a single subapical gland, pubescent, spreading beyond cluster, persistent, clusters contiguous at branch tips. Staminate flowers c.10 per cluster, sessile, stamens 515, anthers 3-locular. Infructescences 15(26) cm pubescent racemes with sparse short 2 cm distal branches (rarely spicate). Fruits in groups of 13 clustered at tips, globular, 12 mm across or shallowly bilobed 10 by 13 mm, woody, densely granular glandular with close or diffuse slender tapering sinuous 5 mm pubescent spines, styles apical, twin, slender, densely shortly pubescent, free to 15 mm long; calyx persistent, 57 mm; pedicel stout, pubescent. Seeds globose, 6 mm, coarsely strongly verrucose, with sarcotesta.
Distribution Moluccas (Aru, Kai, Tanimbar), New Guinea, incl. Bismarck Archipelago, Solomon Islands.
Habitat & Ecology Lowlands to 1000m, secondary forest. Very common.
Note A highly distinctive but variable species from its usually big trilobed leaves and sericeous terminal bud. I have not seen any of the simple-leaved variant Perry mentions.
Macaranga cucullata is similar, but rare.
Macaranga allorobinsonii Whitmore, Gen. Macaranga Prodr. (2008) 72 Type: Robinson 1722 (holo K; iso L), Indonesia, Moluccas, Ambon.
Tree to 2030 m tall. Twigs slender, round, dark, finely furfurescent. Stipules oblong lanceolate, 10 mm, apex acuminate, blackish brown, papery, slightly furfurescent, erect, soon caducous. Leaves: petioles 711 cm, slender, round, glabrous or rarely pubescent, slightly kneed at top; blades elliptic to ovate, 12(28) by 4(16) cm, papery, base rounded to subtruncate with a pair or more of inconspicuous glands at petiole insertion, margin entire, apex acuminate, drying mid slightly olivaceous brown, glabrous or sometimes with spreading white hairs on lower part of midrib below, with close granular glands. Staminate inflorescences slender crowded panicles, c. 10 cm long, branches furfuraceous, secondaries and tertiaries thread-like, secondaries divaricate, peduncle c. 2 cm; 3 branch orders, lowest pair of secondaries opposite, flower clusters towards branch tips, evenly spaced, crowded; bracts tiny, ovate, <1 mm, apex acute, persistent; bracteoles greater than 56 flowered clusters, ovate, c. 3 by 3 mm, apex rounded, (deeply) dentate, nerved, rusty furfuraceous, persistent. Staminate flowers with short pedicel, stamens 4, stamens 4-locular. Infructescences c. 7 cm long, racemes; bracts and bracteoles as staminate flowers, bracts persistent, bracteoles very soon caducous. Fruits sessile, spaced, solitary or in groups of 23, unilocular, or rarely bilobed, round, 4 mm, wall thin, closely granular glandular and sometimes with a few small warts, style near the base 12 mm, shortly feathery, caducous, calyx minute, persistent. Seeds round, surface smooth.
Distribution Sulawesi and Moluccas.
Habitat & Ecology Lowlands to 500m, open or disturbed primary forest.
Notes 1. This species fits the Angustifolia group, distinctive in the basal style, except in having the male bracteoles dentate, larger than the flower clusters and persistent, in which (and the rather crowded inflorescences) it resembles M. polyadenia.
2. Merrill recognised Robinson 1722 as distinct, though reminiscent of his M. robinsonii (now M. myriantha, but to which the epithet alludes), but he declined to name it.
3. Lack & Grimes 1632 differs in having pedicellate flowers with only 2 stamens; it was collected as a `large liane' which I think is erroneous. I tentatively place it here.
Macaranga amentifera Whitmore, Gen. Macaranga Prodr. (2008) 74 Type: Wiakabu & A.M.C. LAE 73557 (holo K), Papua New Guinea, Sepik Prov., Maprik Subprov., Suangagu, 3Ί38S, 143Ί1E.
Small tree to 6m. Twigs slightly flattened, 4 mm diam., smooth, with stipule scars, glabrous, straw-coloured. Stipules elliptic, 15 by 5 mm, apex acuminate, papery, rusty hirsute outside, erect, caducous. Leaves: petioles 16 cm, 3 mm diam., terete, smooth, glabrous except rusty scurfy near top, with short knees at both ends; blades ovate-oblong, 1623 by 1016 cm, thinly leathery, base broadly rounded to truncate, 12 cm peltate with several small glands near petiole insertion, margin recurved in lower part, repand, apex acuminate for 1 cm, drying khaki, glabrous, with or without sparse black granular glands below, 57 nerves at petiole insertion, tertiaries and quaternaries scalariform. Staminate inflorescences amongst leaves, sparsely branched 510 cm panicles, main axis angular 1 mm diam., striate, weakly tomentose or glabrous, peduncle 15 mm, secondary axes short, 12 cm long, lowermost pair opposite, bearing apically dense 10 by 8 mm catkins; axis bracts oblong, 6 by 2 mm, apex acute hirsute, papery, caducous; catkins compound, comprising a dense mass of 6 by 3 mm subcatkins, each subtended by a bract, linear, 3 by 1 mm, apex acute, hirsute outside, subcatkins comprising close oblong overlapping flower clusters of c. 20 flowers, each subtended by a fleshy glabrous bracteole with a 1 by 1 mm slightly cucullate faintly nerved base narrowing fairly sharply into a 1.5 mm erect acumen 0.8 mm wide at its base and protruding beyond cluster, with several small submarginal glands and a few long marginal hairs. Staminate flowers tiny, globose, 0.2 mm diam., sepals 3, free, glabrous, stamens 3, anthers 3-locular, shortly pedicellate. Pistillate flowers, fruits and seeds unknown.
Distribution Papua New Guinea, only known from the type.
Habitat & Ecology Secondary forest, 210 m.
Note A distinctive species of the Brunneo-floccosa group with staminate flowers in dense compound catkins.
Macaranga amissa Airy Shaw, Kew Bull. 21 (1968) 403; Whitmore & Airy Shaw, Kew Bull. 25(1971) 237; Schaeffer, Blumea 19 (1971) 192; Whitmore, Tree Fl.Malaya 2 (1973) 112; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 156; Gard. Bull. Singapore 31 (1978) 53; in Airy Shaw, Kew Bull. 36 (1981) 320; Corner, Ways. Trees Malaya, ed.3 (1988) 299; Whitmore, Gen. Macaranga Prodr. (2008) 75. Type: King's Collector 5267 (holo K; iso CAL?, L), Malaysia, Perak.
Endospermum perakense King ex Hook.f., Fl. Brit. India 5 (1887) 458; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.iv (1912) 35; Ridl., Fl. Malay. Penins. 3 (1924) 306 (non Macaranga perakensis Hook.f., Fl. Brit. India 5 (1887) 447).
Macaranga sp.: Corner, Gard. Bull. Singapore 10 (1939) 299.
Macaranga aff. populifolia auct.non (Miq.) Mόll.Arg.: Corner, Ways. Trees Malaya (1940) 269.
Macaranga sp.: Airy Shaw, Kew Bull. 14 (1960) 396.
Trees to 30 m tall. Twigs slender, to 6 mm diam., terete, coarsely shallowly striate, sometimes with encircling stipules, glabrous. Stipules ovate oblong, 8 by 3 mm, apex acuminate, dark brown, papery, glabrous, erect, caducous. Leaves: petioles 7 cm, 2 mm diam., slender, terete, shallowly striate, glabrous; blades broadly ovate, 814 by 510 cm, (thinly) leathery, drying grey green, base truncate and 1 cm peltate or broadly shallowly cordate, margin recurved, distantly glandular toothed, apex acute to acuminate, below mainly glabrous, slightly glaucous with sparse hairs on main nerves, secondary nerves stout, raised below, same colour as lamina. Staminate inflorescences among and behind leaves, 8 cm, racemes, with a few 3 cm distal alternate catkin-like branches, axis terete, glabrous, peduncle 4 cm; bracts not seen; bracteoles very variable, oblong, 8 by 4 mm, to ovate oblong, 4 by 34 mm, entire or coarsely irregularly lobed or dentate, apex narrowing to obtuse, glabrous, papery, nerved, blackish brown, longer than cluster, erect, subpersistent. Staminate flowers in clusters of 5, oblong, 0.5 mm, glabrous; pedicel 0.5 mm; sepals 3, free, oblong; stamens 2, anthers 4-locular. Infructescences stout 10 cm spikes, axis 2 mm, terete, rugose, peduncle to 3 cm. Fruits well spaced, single, shallowly bilobed, slightly compressed, 5 by 9 by 4 mm, blackish granular glandular, woody, stigmas tiny, apical, persistent, pedicel 23 mm, stout, glabrous; sepals 3, recurved, persistent. Seeds with a thin sarcotesta, globose, testa with flat warts.
Distribution Malay Peninsula (rare but gregarious), Sumatra (Palembang, Bangka), Borneo (Sabah).
Habitat & Ecology Primary or secondary usually swampy forest.
Macaranga amplifolia Merr., Philipp. J. Sc. 7 (1912) 392, Enum. Philipp. Flow. Pl. (1923) 440; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 317; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 33; Whitmore, Gen. Macaranga Prodr. (2008) 76. Type: Ramos BS 13513 (holo PNH, lost; iso BO, K, L), Philippines, Luzon, Laguna, San Antonio.
Tree 10m. Twigs stout, terete, c. 1 cm, rufous velvety, at first densely so. Stipules elliptic, 45 by 5mm, apex acuminate, blackish, papery, lanate with long silky hairs, glabrescent. Leaves: petioles to 16 cm, slender, terete, finely striate, glabrous or with scattered white hairs; blades big, deltoid, 1735 by 920 cm, thickly papery, base broadly truncate, very slightly rounded, margin with distant small glands, apex acute, glabrous, granular glandular below. Inflorescences amongst and behind leaves, crowded panicles to 28 cm, 3(4) orders of somewhat flattened furfurescent axes, secondaries divaricate, spaced groups of 34 flower clusters; male bracteoles persistent, tiny, oblong, 2 by 1mm, apex blunt, distal end coarsely bluntly toothed, furfuraceous. Staminate flowers 4 per cluster, pedicellate, 0.5 mm long; sepals fused; stamens 2, anthers 4-locular. Fruits dicoccous, 4 mm cocci, glandular, unarmed;, pedicel 2 mm; calyx persistent. Seeds round, finely verrucose.
Distribution Philippines (Luzon, Leyte).
Note Fruit description after Merrill. Except for the glabrous leaves this is similar to the northern form of M. noblei.
Macaranga anceps Airy Shaw, Kew Bull. 19 (1965) 324; Whitmore in Airy Shaw, Kew Bull. 36 (1981) 319; Gen. Macaranga Prodr. (2008) 76. Type: Achmad 1445 (holo K, iso L), Sumatra Simalur [Simeulue] Island.
subsp. anceps
subsp. anceps: Whitmore, Gen. Macaranga Prodr. (2008) 76. Type: as species.
Small tree to 15 m. Twigs slender 3 mm diam., terete, smooth, glabrous, warm brown, becoming rugose with age. Stipules and bud scales elliptic, 3(5) by 1 mm, brown, leathery, stiff, glabrous, erect, crowded at apex, soon caducous. Leaves: petioles 2.5 cm, slender, 1 mm diam., terete, striate, glabrous, kneed at top; blades elliptic, 915 by 35.5 cm, thickly papery, drying brown to grey green, base usually long cuneate rarely rounded, with 2 miniscule gland-bearing auricles at petiole insertion, margin weakly sinuous, apex abruptly 2 cm acuminate glabrous, below without puncticulations but occasionally with sparse chocolate granular glands. Staminate inflorescences spikes to 2 cm, squarish in section, finely pubescent, with flower clusters becoming 4 mm spaced, peduncle 05 mm; bracteoles triangular, 0.5 by 0.4 mm, apex acuminate, spreading, entire, glabrous, persistent, smaller than cluster. Staminate flowers 23 per cluster, 1 mm diam.; sepals oblong, 1 by 1 mm hooded, apex obtuse, gland tipped, granular glandular outside; stamens 10, filamens 0.5 mm long, anthers 4-locular. Pistillate inflorescences sinuous 10 cm spikes, axis glabrous (rarely furfuraceous), strongly flattened, 2 mm wide, striate; peduncle 7 cm; flowers single, irregularly spaced near tip, often subopposite; bracts not seen; bracteoles elliptic, 1.5 mm, stiff, erect, furfurescent on both surfaces, most persisting. Pistillate flowers not seen. Young fruits: pedicel 3 mm, pubescent; sepals elliptic, 2 mm, pubescent; ovary 3 mm, tawny pubescent and densely softly spiny, with twin 15 mm slender sinuous smooth apical styles. Mature fruits with 2 globular lobes, each 6 mm diam., tawny pubescent, spaced conico-muricate from the bases of earlier spines, thinly wood; pedicel to 5 mm; sepals persistent, recurved; styles caducous. Seeds spherical, 5 mm diam., smooth, shiny, with a thin sarcotesta.
Distribution Sumatra (including P.Simeulue), Borneo (Sarawak).
Habitat & Ecology Primary forest.
subsp. puncticulata Whitmore
subsp. puncticulata Whitmore, Kew Bull. 29 (1974) 450, in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 154; Gen. Macaranga Prodr. (2008) 77. Type:, Sibat ak Luang S 24517 (holo K, iso L), Borneo, Sarawak, 4th Division, Bintulu, Nyabau Catchment.
Distribution Peninsular Malaysia (Trengganu), Borneo (Sarawak, Brunei).
Note Differs in Petiole stouter, 2 mm diam; blades below densely covered with volcano-like puncticulations, shiny and leathery.
Macaranga andamanica Kurz, For. Fl. Brit. Burma 2 (1877) 389; Pax & K.Hoffm. in Engl., Pflanzenr. VI.147.vii (1914) 365; Parkinson, Forest Fl. Andaman Is. (1923) 238; Airy Shaw, Kew Bull. 19 (1965) 318; Kew Bull. 21 (1968) 405; Kew Bull. 26 (1972) 287; Whitmore, Gard. Bull. Singapore 31 (1978) 53, Tree Fl. Malaya 2 (1973) 112; Chiu, Guihaia 2(1982)152; Chakrab. & N.P.Balakr., J. Econ. Taxon. Bot., Addit. Ser. 9 (1992) 78; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 363; Whitmore, Gen. Macaranga Prodr. (2008) 77. Type: not indicated.
Macaranga brandisii King ex Hook.f., Fl. Brit. India 5 (1887) 453; Brandis, Indian Trees (1906) 592; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914)365; Ridl., Fl. Malay. Penins. 5 (1925) 333.
Morinda esquirolii Lιv., Fl. Kouy-Tchιou (1915) 368. Macaranga esquirolii (Lιv.) Rehder, J. Arnold Arbor.18 (1937) 214; Croizat, J. Arnold Arbor. 23 (1942) 214, passim.
Macaranga kampotensis Gagnep., Bull. Soc. Bot. France 69 (1922, published 1923) 702, Lecomte, Fl. Indo-Chine 5(1926) 447.
Macaranga bracteata Merr., Lingnan Sci. J. 6 (1928) 281; Chiu, Guihaia 2 (1982) 251; Airy Shaw, Kew Bull. 21 (1968) 406, passim.
Macaranga rosuliflora Croizat, J. Arnold Arbor. 23 (1942) 51; Airy Shaw, Kew Bull. 21 (1968) 406, passim; Chiu, Guihaia 2 (1982) 149.
Macaranga trigonostemonoides Croizat, J. Arnold. Arbor. 23 (1942) 51; Airy Shaw, Kew Bull. 21 (1968) 405; Chiu, Guihaia 2 (1982) 150.
Treelet to 5 m. Twigs slender, 3 mm diam., terete, striate, glabrous. Stipules and bud scales elliptic, 8 mm, with a caudate or subulate tip, chocolate brown, stiff but somewhat flexuous, with sparse spreading white whiskers, crowded at twig tip, erect, soon caducous. Leaves: petioles 1.53 cm, very slender, terete, glabrous or with pale crisped and spreading hairs; blades elliptic, 711 by 25cm, papery, drying dark to mid brown or rarely khaki green, base rounded to long cuneate, minutely auriculate, with 2 glands near petiole insertion, margin weakly crenate and glandular dentate, apex (abruptly) acuminate, usually glabrous sometimes pubescent on lower part of midrib below, sparse dark granular glands. Staminate inflorescences never forming witches' brooms, delicate thread-like spikes, 5(12) cm, glabrous or with pale crisped and spreading hairs (like petiole), terete; peduncle short; flower clusters tiny, 11.5 mm across, regularly, c. 3 mm spaced, like beads on a slightly zigzag string; bracteoles persistent, ovate 1 mm, apex acute, strongly hooded and enclosing the flower cluster, outside finely furfuraceous, entire. Staminate flowers in clusters of c. 3, developing in sequence, the mature one exserted beyond bracteole on a pubescent 0.5 mm persistent pedicel; sepals 3, granular glandular; stamens c. 15, anthers 4-locular. Infructescences 36 cm spikes with terminal fruits; axis slender, terete, glabrous or pubescent. Fruits: a single developing subtended by 2 opposite variable sessile leafy bracts, elliptic, 1 cm long, or ovate, to 2.5 by 1cm, apex acute; fruits strongly dicoccous, 5 by 11 by 5mm, woody, closely black-granular glandular, with sparse long sinuous caducous spines; pedicel 15 mm stout pubescent; sepals free, persistent; twin styles 1 cm thread-like, smooth.
Distribution India (Andamans), Myanmar (Amherst, Tenasserim), China (Yunnan, Guangdong, Kweichou, Guangzou, Hainan), North Vietnam, Cambodia, Thailand (Peninsula), Peninsular Malaysia.
Habitat & Ecology Lowland and lower montane primary forests (always?).
Notes 1. This is a widespread and highly variable species as Airy Shaw (1965: 318) has pointed out. Its distinctive characters are the solitary fruit terminal on a slender spike and subtended by (very variable) leafy bracts. This fruit is strongly bilobed and black granular glandular with evanescent spines. The staminate spike is delicate, with evenly spaced small flower clusters. The chocolate brown stipules have sparse spreading white whiskers. The plant is usually glabrous, but may bear pale crisped and spreading hairs on petiole, inflorescence axis and midrib below.
2. Macaranga lowii, which is also polymorphic, is very similar in habit, but has stiff chestnut brown stipules, which are glabrous (except Tsang 376, Tsiang 2223), pistillate flowers and fruits not terminal on the spikes and stouter staminate spikes with larger flower clusters.
3. Macaranga digyna, a Sri Lankan endemic, is closely related. It has the pistillate parts like M. lowii, the staminate delicate like M. andamanica and glabrous to furfuraceous sinuate stipules like neither.
4. I am quite unable to maintain M. trigonostemonoides. The characters used by Airy Shaw (1968: 405) to distinguish it from M. andamanica are actually contradicted by its type.
5. From its description, M. kampotensis from southern Cambodia indubitably comes here, and M. esquirolii probably likewise.
Macaranga angulata S.J.Davies, Kew Bull. 54 (1999) 150; Harvard Pap. Bot. 6 (2001) 385. Type: Clemens 40736 (holotype: A; isotypes: K, L), Malaysia, Sabah, Gunung Kinabalu.
Small tree to 10 m tall with smooth grey-brown bark; twigs 48 mm in diam., glabrous or rarely with sparse minute hairs, hollow, housing ants, not glaucous. Stipules deltate, 610 by 610 mm, recurved, succulent, producing food-bodies on abaxial surface, up to 4 pairs present on the shoots. Leaves: petioles terete, 820 cm long, glabrous; blades broadly to very broadly ovate, 1430 by 1122 cm, deeply trilobed, sometimes with two additional basal leaf lobes or cusps, lateral lobes narrow and spreading, dissected to 1/22/3 of the leaf length, 13 cm peltate, base very broadly rounded to truncate, margin entire, sometimes with small protruding, flattened nectaries, apices acute to finely acuminate, adaxial surface glabrous, abaxial surface glabrous or rarely with very tiny scattered hairs on veins, not glaucous; 1° venation palmate with 89 prominent veins, 2° venation scalariform, looping near the leaf margins; young leaves red-brown, finely ferrugineus pubescent, apices acute to acuminate. Staminate inflorescences paniculate, erect, 1113 by 710 cm, glabrous, 34 axis orders, main axis flattened, first pair of secondary branches opposite with accessory branches; bracts deltate, 610 by 34 mm, caducous, margin entire, apex acute, glabrous or with scattered hairs; flower clusters with 2030 flowers, spirally arranged and evenly spaced along inflorescence branches; bracteoles ovate, 35 by 34 mm, enclosing flower clusters and often strongly reflexed after anthesis, margin entire, apex acute to shortly acuminate, scattered to densely covered with minute red-brown hairs and yellow granular glands. Staminate flowers c. 1.0 mm long, sessile; sepals fused, splitting irregularly, apex densely covered in minute red-brown hairs; stamens one; anthers 3-locular. Pistillate inflorescences particulate, erect, 410 by 48 cm, glabrous or rarely with minute scattered hairs, up to 2 axis orders; bracts caducous. Pistillate flowers 2.53 by c. 2. mm, solitary in bracteole axils; calyx urceolate, c. 1.5 mm long, apex with fine red-brown hairs and yellow granular glands, persistent, splitting irregularly as ovary expands; ovary 45-carpellate; styles c. 5 mm long, fused at base, free and spreading from 1/23/4 of the length, persisting to form a prominent 34 mm long crown at fruit apex; stigma not dissected. Fruits 56 by 911 mm, subglobose, sessile, one discrete glandular patch on each carpel wall developing into a slender horn-like process 46 mm long, covered in yellow, sticky exudate. Seeds subtriangular-ovoid, c. 4 mm in diam., black, with shallow coarse grooves and a small cruciform scar, encased in a fleshy pink-red aril.
Distribution Borneo (Brunei, Sabah, Sarawak).
Habitat & Ecology Early successional tree in lower montane forest from 800 to 1500 m in both disturbed and primary forests. This species is known from Gunung Kinabalu south along the mountains into Brunei and the Kelabit Highlands in northern Sarawak. It probably occurs further south in Sarawak and in the mountains of east Kalimantan although it is yet to be collected from there.
Notes 1. This is one of the few ant-inhabited species of Macaranga that grow at over 1000 m altitude. It differs from related species in having deeply dissected 35 lobed leaves which appear quite angular because of the spreading lobes and the almost truncate leaf base, the petiole inserted only a short distance from the leaf margin, and the acute to shortly acuminate staminate bracteoles which have entire margins.
2. Davies (1999) mentioned that several specimens were slightly aberrant in having particularly variable leaves; recent more detailed field work suggests that all specimens belong to M. angulata.
Macaranga angustifolia Lauterb. & K.Schum. in K.Schum. & Lauterb., Fl. Schutzgeb. Sόdsee (1900) 398; Nachtr. (1905) 297; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 379; L.M. Perry, J. Arnold Arb. 34 (1953) 230; Airy Shaw Kew Bull.23 (1969) 110 passim; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)139; Gen. Macaranga Prodr. (2008) 79. Type: Lauterbach 2228 (holo: B, lost; iso K mistakenly numbered as 228), Papua New Guinea, Sepik area, Nuru River.
Small tree 8(16) m. Twigs slender, 23 mm diam., slightly angled and ridged, glabrous, blackish brown. Stipules elliptic, 8 mm, apex acuminate, blackish brown, papery, glabrous, erect, soon caducous. Leaves: petioles 1.22 cm, slender, 11.5 mm diam., slightly flattened, usually somewhat channelled above, glabrous, weakly kneed at top; blades elliptic, 16 by 36cm, thinly leathery, drying mid brown, base cuneate, 2 elongate usually narrow marginal glands near petiole insertion, margin usually slightly repand, often with small inconspicuous distant protruding glands, recurved slightly but sometimes more strongly near base, apex abruptly 5 mm acuminate, glabrous, closely finely sometimes dark granular glandular, basal secondary nerves (sub)marginal. Staminate inflorescences amongst leaves, fairly dense, narrow, 913 cm, glabrous or weakly furfuraceous racemes; branches to 4 cm, ascending, some arising from a flower cluster; peduncle 13 cm, main axis slender, 1 mm diam., flattened, lower branches alternate or (sub)opposite, thread-like with evenly 23 mm spaced flower clusters; bracteoles conspicuous, variable (elliptic to) ovate, 3 by 2 mm, apex acute to acuminate, or 2 by 2 mm, apex blunt, with 2 very large 1 mm diam. adaxial patellar glands, margin dentate, glabrous, fleshy, spreading or reflexed, a few persistent, larger than cluster. Staminate flowers in clusters of c. 20, developing sequentially, shortly pedicellate; stamens c. 20, anther 4-locular. Infructescences as staminate but shorter, usually to 6 cm, and often spikes; bracteoles soon caducous. Fruits in groups of c. 4, tiny, globose, 3 mm diam., thinly woody, densely usually blackish granular glandular, with 12 short persistent slender (? brittle) horns; pedicel slender, 5 mm, glabrous; calyx caducous, style plumose, bifid, basal, soon caducous. Seeds globose, finely verrucose, with sarcotesta.
Distribution New Guinea (throughout, widespread).
Habitat & Ecology Open primary forest, riverbanks, and secondary forest. Lowlands to 1350 m. Common.
Note The conspicuous bracteoles are highly diagnostic, though soon caducous in the pistillate. In leaf this closely resembles M. pleiostemon, but the leaf base glands are usually narrowly elongate.
Macaranga ashtonii S.J.Davies, Harvard Pap. Bot. 6 (2001) 386. Type: Davies 99309 (holotype: SAR: isotypes: A, K, KEP, L, SING), Malaysia, Sarawak, near Puncak Borneo, Padawan area, 1°08'N, 110°15'E.
Large submontane tree to 25(30) m tall and 30 cm dbh, with big pendulous leaves and a red-brown appearance; twigs 1020 mm diameter, strongly angular when dry, terete when fresh, with prominent whitish lenticels, densely ferrugineus pubescent, particularly towards the apex, sometimes slightly glaucous, solid, exuding clear to pale reddish latex when cut, not housing ants. Bark smooth, pale brown, hoop marked. Stipules deltate, (10)1525 by (8)1020 mm, apex acute, initially erect but often spreading to recurved, not glaucous, coriaceous, densely covered in reddish golden hairs, the hairs very dense towards the base but sparse towards the margins, the pair almost completely encircling the twig, 13 pairs present on the shoots. Leaves: petioles terete but drying slightly flattened, stout, (12)1535 cm long, 48 mm in diam., greyish, densely pubescent with erect silvery hairs; blades broadly ovate, large to very large, 2560 by 1540 cm, trilobed, dissected to c. 1/41/2 of the leaf length, central lobe broadly deltate and up to 20 cm wide at the base. lateral lobes erect, leaf lobes up to 8 cm wide at the base, 49 cm peltate, base broadly rounded, margin entire, with numerous small conical nectaries, leaf apices sharply acute, adaxial surface glabrous or with fine ferrugineus hairs along the veins, shiny green with reddish venation when fresh, abaxial surface densely pubescent with erect silvery hairs and minute ferrugineus hairs along the veins, greyish to slightly glaucous, without surface patellary nectaries; 1° venation palmate with (7)89 prominent veins, 2° venation prominent and densely scalariform, looping and terminating at the margin in fine conical nectaries, 3°4° venation prominently scalariform; young leaves pale pinkish brown with the margin turning green first, adaxial surface densely covered in minute ferrugineus hairs, abaxial surface densely covered in erect silvery hairs and small yellow granular glands. Staminate inflorescences paniculate, erect, c. 15 by 10 cm (not fully mature), densely ferrugineus pubescent, up to 4 axis orders, main axis flattened, first pair of secondary branches ±opposite with accessory branches; bracts deltate, 511 by 36 nun, margin entire, apex acute, sparsely to densely ferrugineus pubescent on abaxial surface and margin, adaxial surface ± glabrous; flower clusters with c. 10 flowers, decussately arranged and clustered on ultimate inflorescence branches; bracteoles (immature) broadly ovate, 16 by 34 mm, enclosing flower clusters, margin finely and shallowly pectinate to almost entire, apex acute ending in a tiny conical gland, abaxial surface densely pubescent with fine red-brown hairs and scattered with yellow granular glands, adaxial surface mostly glabrous with a dense patch of ferrugineus hairs at the base. Staminate flowers c. 1 mm long, pedicellate; sepals free, apex scattered with minute red-brown hairs; stamens 3 or 4; anthers 3(4)-locular. Pistillate inflorescences paniculate, erect, 1015 by 612 cm, stout, densely golden ferrugineus pubescent, up to 3 axis orders, secondary branches ± opposite; bracts deltate, c. 7 by c. 4 mm, densely golden ferrugineus pubescent, caducous. Pistillate flowers 46 mm long, solitary; calyx urceolate, c. 4 mm long, very finely golden ferrugineus pubescent, apex with yellow granular glands, persistent, splitting irregularly as ovary expands; ovary 45-carpellate, c. 4 mm long; styles c. 1.5 mm long, fused at base, free and spreading from c. 1/2 of the length, persisting to form a prominent 23 mm long crown at fruit apex; stigma not dissected. Fruits (immature) 1012 by 68 mm, subglobose, compressed, shortly pedicellate, one discrete glandular patch covering most of each carpel wall but the sutures not glandular, at maturity yellow sticky exudate covering most of the fruit but not raised; pedicel 36 mm long, finely ferrugineus pubescent. Seeds (immature) subtriangular-ovoid, c. 4.5 mm in diam., brown-black, with shallow coarse grooves and a small cruciform scar at the columella attachment point, encased in a fleshy aril (colour not seen).
Distribution Borneo (Sabah, Sarawak).
Habitat & Ecology This species has been collected from four locations in submontane forest in Borneo from 800 to 1300 m elevation. Macaranga ashtonii is a large tree in primary and secondary submontane forest. Abundant regeneration occurs in logged forest and along roadsides.
Notes 1. This species is named in honor of Professor Peter S. Ashton of Harvard University for his significant contribution to the ecology and systematics of Bornean trees.
2. Macaranga ashtonii is a striking tree in submontane forest. The collecting localities stretch from Gunung Trus Madi in Sabah in the north, through the Kelabit Highlands of northern Sarawak, to the area of Gunung Penrissen near Padawan in the south. Davies (2001) suspects that it occurs throughout the submonane forests of central Borneo, although it was not found during his recent trip to the summit of Gunung Murud in the Kelabit Highlands.
3. It appears to be most closely related to another recently discovered Bornean endemic, Macaranga glandibracteolata. With that species it shares finely pectinate staminate bracteoles terminated by a small conical gland, 34-thecate anthers, 45-carpellate fruits without prominent horn-like processes and prominent 3°4° venation on abaxial leaf surfaces. Macaranga ashtonii differs from M. glandibracteolata in having twigs that are densely ferrugineus hairy, not ant inhabited, not or only slightly glaucous, and strongly angular when dry. Macaranga ashtonii also has less deeply pectinate staminate bracteoles, and the glandular patches cover almost the entire fruit wall. Furthermore, M. ashtonii is a submontane tree and M. glandibracteolata is restricted to lowland forest.
Macaranga astrolabica Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 343; L.M.Perry, J. Arnold Arb. 34 (1953) 231; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 161; Gen. Macaranga Prodr. (2008) 81. Type: F.H. Brown 165 (not seen), Papua New Guinea, Madang Prov., Astrolabe Bay.
Twigs slender, glabrous. Stipules narrowly triangular, 2 mm, soon caducous. Leaves: petioles 24 cm, glabrous; blades elliptic, 817 by 3.55.5 cm, subcoriaceous, base rounded narrowly cordate, with 13 glands near petiole insertion, margin entire or wavy, long acuminate, granular-glandular, glabrous, with 3 basal nerves and 67 pairs of secondaries, reticulations visible below. Staminate inflorescences panicles 48 cm long, narrow, sparsely branched and flowered; peduncle glabrous; branches puberulous; bracts ovate triangular, aepx acute, puberulous without glands; bracteoles elliptic, 23 mm long, narrowed at base, entire or repando-lobulate, apex acute, glabrous or puberulous, with patelliform glands inside, spreading. Staminate flowers 1 mm diam.; sepals 2, ovate, concave, granular-glandular outside at tip; stamens 2, anthers 4-locular. Pistillate inflorescences, flowers, fruits and seeds unknown.
Distribution New Guinea (only known from type).
Notes 1. Only known from the type collection. Not seen by me. The description is paraphrased from Perry's translation, amended against the original Latin.
2. From the description this species has no very striking features. However, it keys out with the Dioica group. I therefore place it there provisionally.
Macaranga auctoris Whitmore, Gen. Macaranga Prodr. (2008) 82 Type: Soejarto & Fernando 7445 (holo L), Philippines, Palawan, Taytay, Ibangley, Pagdanan Range, road Taytay Roxas, 10o35' N, 119o33' E
Treelet 5 m. Twigs stout 15 mm diam., strongly angled, coarsely striate, black, sparsely rusty furfuraceous. Stipules not seen. Leaves: petioles 45 cm, stout, 8 mm diam. at base, flattened above, but basally round, glabrous, blackish brown, kneed at top; blades big, ovate-elliptic, 52 by 28 cm, broadest 12 cm from base, leathery, base very broadly rounded, 5 cm peltate, margins recurved, irregularly shallowly crenate, apex tapering to acute, drying grey, below paler grey with the main nerves brown, glabrous, with below irregularly patchy sparse dark brown granular glands, above glabrous, 5 main and numerous lesser nerves from petiole insertion, secondaries 12 pairs, distant, slightly curved, parallel, tertiaries and quaternaries scalariform. Staminate inflorescences amongst leaves as a dense 8 cm diam. cluster near twig tip, panicles (?); peduncle 1cm, flattened 3 mm across, striate, glabrous; lowest branches subopposite; ultimate branches to 1 cm arising from a flower cluster; clusters in a tight spiral, looser distally; bracteoles ovate, 57 by 45 mm, including deeply coarsely fimbriate margins and caudate tip, strongly concave, papery, outside with scattered golden granular glands and sparse short hairs, faintly nerved, much larger than clusters, overlapping, persistent. Staminate flowers 5 per cluster, shortly pedicellate; sepals free, outside with granular glands; stamens 2, anthers 4-locular. Pistillate inflorescences, flowers, fruits and seeds unknown.
Distribution Philippines (Palawan, known from type only).
Habitat & Ecology Logged lowland rain forest, 5070 m.
Notes 1. A Macaranga for the connoisseur, with its big grey peltate leaves and crowded inflorescences bearing purple red flowers, this must be a striking treelet. I name this species in the same spirit that certain choice vintage wines are labelled `directors' reserve'. It must be an easy plant to spot, though I have not myself seen it on either of my two visits to Palawan.
2. The greyness of the leaves is not a wax: the Kuhlman test failed. I do not know where to place this species. The big peltate leaves echo the Mappa group. The rather large fimbriate, nerved bracteoles are reminiscent of M. congestiflora (Bicolor group), which also has congested inflorescences, and is also confined to Palawan.
Macaranga baccaureifolia Airy Shaw, Kew Bull. 19 (1965) 316; Whitmore, Tree Fl.Malaya 2 (1973) 112, in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 154; Gen. Macaranga Prodr. (2008) 83. Type: Corner SFN 29287 (holo K; iso SING), Peninsular Malaysia, Johore, Sungai Berassau, Mawai Jemalung Road.
Small tree to 12 m. Twigs slender 3 mm, angled, smooth, rusty furfuraceous. Stipules 6 mm, subulate, widened at base, erect, slightly pubescent. Leaves: petioles 811 cm, slender, 2 mm diam., weakly angled and striate, rusty furfuraceous, kneed at both ends; blades broadly oblong ovate (9)21 by (5)12 cm, papery, base broadly rounded and cordate (or rarely 2mm peltate), minutely auriculate the lobes touching to slightly overlapping, eglandular, margin undulate, apex acuminate, narrowing sometimes abruptly to a 2 cm tip, warm brown, glabrous to (sub)pubescent on main nerves below and granular glandular. Staminate inflorescences spikes to 5 cm, terete, pubescent; peduncle absent; bracts not seen; flower clusters more or less contiguous; bracteoles spreading, ovate to 2 by 2 mm, shouldered or with 2 lateral cusps, apex acute, granular glandular without. Staminate flowers developing in sequence, sessile, globular, 1 mm diam.; sepals granular glandular and pubescent outside; stamens c. 10, anthers 4-locular. Infructescences crowded amongst apical leaves, spikes to 9 cm, axis flattened, pubescent, with a single terminal fruit subtended by 2 bracts, latter leafy, orbicular, 2.5 cm across, apex acute to rounded, pinnately nerved, abaxially densely and adaxially sparsely granular glandular, and with sparse abaxial setose hairs. Fruits strongly dicoccous, thickly woody, lobes globular, 1 cm diam., densely shortly tomentose; pedicel 1 mm, tomentose; sepals persistent; styles to 1.5 cm, caducous. Seeds round, 6 mm diam., smooth, without sarcotesta.
Distribution Peninsular Malaysia (Johore), Sumatra, Borneo (Sabah).
Habitat & Ecology Primary forest, in Malaya freshwater tidal swamp forest.
Note Corner (Freshwater swampfor. S. Johore Singapore, 1978) notes that in Johore this is gregarious and loses its leaves when the annual flooding recedes every February.
Macaranga balabacensis Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 368; Merr., Enum. Philipp. Flow. Pl. (1923) 440; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 33; Whitmore, Gen. Macaranga Prodr. (2008) 83. Type: Mangubat 424 (isotype BO), Philippines, Mindoro, Balabac.
Similar to M. hispida but conspicuously differing in Leaves (3)10 mm peltate, base broadly 1015 mm cordate, or truncate, upper side with c. 6 glands between petiole insertion and blade. Staminate bracteoles 57 mm long, petiolate, ovate-lanceolate, apex acuminate, deeply narrowly dentate, teeth tipped with conspicuous adaxial patellar glands, villous, granular glandular on both surfaces.
Distribution Philippines (Mindoro: Balabac; known from the type only).
Macaranga bancana (Miq.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 990. S.J.Davies, Harvard Pap. Bot. 6 (2001) 387; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 364. Pachystemon bancanus Miq., Fl. Ned. Ind., Eerste Bijv. (1861) 462. Type: Teijsmann s.n. (holotype: L; isotype: BO), Indonesia, Sumatra, Insula Bangka prope Djebus.
Macaranga tenuifolia Mόll.Arg. in DC., Prodr. 15, 2 (1866) 990. Type: Teijsmann s.n. (G-DC, not seen), Indonesia, Sumatra, Bangka.
Macaranga triloba auct. non. (Thunb.) Mόll.Arg.; Merr., Enum. Philipp. Flow. Pl. (1923) 448; Ridl., Fl. Malay. Penins. 3 (1924) 298; Corner, Ways. Trees Malaya (1940) 304; Whitmore, Tree Fl. Malaya 2 (1973) 107; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 150; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 35.
Macaranga calcifuga auct. non. (Whitmore) R.I.Milne: Milne, Kew Bull. 49 (1994) 453, p.p.
Abundant small tree to 23 m tall, but reproductive from only 23 m tall; twigs 49 mm in diam., glabrous or rarely with scattered minute ferrugineus hairs towards the apex, not glaucous, hollow, housing ants. Bark pale-brown to whitish, mottled, smooth with small lenticels, hoop-marked. Stipules broadly ovate, 68 by 69 mm, red-brown when fresh, usually finely furfuraceous on the base of the adaxial surface when young and becoming glabrous, the pair completely encircling the twig, succulent, recurved, producing food-bodies on the abaxial surface, usually 24 pairs present on the shoots. Leaves: petioles terete, slender, 824 cm long, not glaucous, glabrous; blades broadly ovate, 1632 by 1225 cm, trilobed to rarely just tricusped, variably dissected to 1/81/2 of the leaf length, central lobe broad sometimes constricted near the base, lateral lobes ascending and 2.55(7) cm wide, rather shallowly peltate, petiole insertion 14 cm from margin, base broadly rounded, margin entire, apices narrowly acute, adaxial surface glabrous or scattered with fine ferrugineus hairs along the veins when young, abaxial surface glabrous with the veins sparsely to evenly covered in minute crisped silvery hairs; 1° venation palmate with 78 prominent veins, 2° venation scalariform and rather closely spaced usually with 1416 pairs of veins arising from the midrib, curving upwards slightly near the leaf margins and running into the margin and terminating in tiny (c. 0.3 mm) narrow conical nectaries; young leaves reddish-brown, densely minutely ferrugineus hairy, soon becoming glabrous. Staminate inflorescences furfuraceous, reddish-brown, paniculate, erect, 1020 by 515 cm, densely ferrugineus tomentose, particularly dense towards the apex and becoming more scattered to rarely almost glabrous at the base, with the hairs also slightly longer towards the base, up to 4 axis orders with the distal branches characteristically looping and convoluted, main axis flattened, first pair of secondary branches opposite with accessory branches; bracts ovate, 36 by 23 mm, margin entire, apex acute, finely ferrugineus pubes cent on both surfaces and the margin, caducous; flower clusters with 1525(30) flowers, spirally arranged and scattered along ultimate inflorescence branches: bracteoles small, broadly ovate to transversely elliptical, 1.52.5 by 24 mm, enclosing flower clusters, margin entire, apex rounded or with a short broad acute tip, very densely covered with minute ferrugineus hairs and yellow granular glands. Staminate flowers c. 0.7 mm long, sessile; sepals fused, splitting irregularly to c. 1/3, apex densely covered in minute red-brown hairs; stamens 1, anthers 3-locular. Pistillate inflorescences paniculate, erect, 311 by 37 cm, stout, scattered with short ferrugineus hairs towards the base and more dense towards the apex, up to 3 axis orders, secondary branches opposite and usually with accessory branches; bracts elliptic, 58 by c. 3 mm, apex acute, abaxial surface scattered with minute ferrugineus hairs and tiny yellow granular glands particularly towards the base, adaxial surface scattered with ferrugineus hairs, caducous. Pistillate flowers c. 2 mm long, solitary in bract axils; calyx urceolate, 1.01.5 mm long, densely ferrugineus pubescent and scattered with yellow granular glands, persistent, splitting irregularly as ovary expands; ovary 4-(sometimes 5-)carpellate, c. 1.5 mm long: styles c. 1 mm long, fused at base, free and spreading from c. 1/2 of the length, persisting to form a prominent 23 mm long crown at fruit apex; stigma not dissected. Fruits 57 by 1011 mm, bright green and slightly reddish on the sutures when fresh, subglobose, compressed, sessile, with scattered ferrugineus hairs on the sutures and at the base of the styles, with one discrete glandular patch on each carpel wall developing into a short pointed swelling c. 12 mm long, covered in yellow-green, sticky exudate. Seeds subtriangular-ovoid, c. 4 mm in diam., black, with shallow coarse grooves and a small cruciform scar at the columella attachment point, encased in a fleshy bright pinkish to orange-red aril.
Distribution Peninsular Thailand, Malay Peninsula, Sumatra (Bangka, Lingga Archipelago, N of Palembang), Borneo (Brunei, Sarawak and Sabah, in latter west of Gunung Kinabalu and the Crocker Range and Sarawak).
Habitat & Ecology This is one of the most common and widespread species in section Pachystemon. Macaranga bancana occurs in a broad range of habitats in the lowlands below about 700 m. It is common on both clay and sandy soils, although clearly prefers the latter, which may in part be due to the water status of those soils, as it is also quite a common colonist of wet gullies along roadsides, alluvial forest areas, and the margins of degraded peat swamp forests.
Notes 1. Davies (2001) has not seen the Teijsmann collection that is the type of M. tenuifolia, however it seems most likely to be M. bancana, as all Macaranga section Pachystemon collections of Teijsmann from Bangka that Davies (2001) has seen are M. bancana and the description sounds right for that species.
2. Macaranga bancana was previously mistaken for M. triloba. However, it differs greatly from M. triloba in having small rounded bracteoles, hollow ant-inhabited twigs, very shortly horned fruits, and much less prominently dentate leaf margins. Macaranga bancana is more closely related to M. velutina and M. indistincta. Both M. velutina and M. indistincta differ from M. bancana in having densely pubescent lower leaf surfaces. In addition, M. velutina often has bluish glaucous pubescent twigs, silvery hairs on the inflorescence axes, and bluish green fruits. Macaranga indistincta differs from M. bancana in having long horned fruits and pectinately dissected staminate bracteoles.
Macaranga barkeriana Whitmore, Gen. Macaranga Prodr. (2008) 84, Fig. 2. Type: C. Barker CB23 (holo K; iso BO, CANB, FRE, GH, L, MAN, SING).
Similar to M. hispida, distinguished by: Tree to 1015m. Twigs very coarsely densely hispid with spreading pale golden 23 mm tufted hairs. Stipules larger, commonly 13 cm long, ultimately before falling gigantic, 24 by 8 cm; apical bud strongly gibbous, to 3 cm across. Leaves: petioles flattened above, 5 mm across, hispid as twigs; blades mostly relatively broader 1736 by 12 30 cm, sometimes shallowly trilobed, mostly (sub)glabrous above except for nerves, below granular glands golden. Staminate inflorescences hispid with 4 axis orders, main axis stouter, 4 mm across near base, basal and some other branches opposite; bracteoles larger, 5 by 2 mm, apical half wholly covered by c. 14 patellar glands. Staminate flowers: anthers 3- or 4-locular. Infructescences racemose, axes hirsute as twigs, with a pair of opposite divaricate 8 cm branches, subtending bract larger, 50 by 8 mm; peduncle 3 cm. Ripe fruit unknown?
Distribution Indonesian New Guinea (Papua: Mimiku Regency near Tembagapura).
Habitat & Ecology Lower montane rain forest 15401590m.
Note This is a coarsely hirsute relative of M. hispida, bigger in its parts and with more copiously branching inflorescences, but otherwise very similar. It is named for the collector, Christine Barker, botanist at Kew.
Macaranga beccariana Merr., Webbia 7 (1950) 315; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 145; S.J.Davies, Harvard Pap. Bot. 6 (2001) 391. Lectotype (designated by S.J.Davies, 2001): Beccarii PB 1257 (lectotype: A; isolectotype; K), Malaysia, Sarawak, Kuching.
Macaranga hypoleuca (Rchb.f. & Zoll.) Mόll.Arg. var. borneensis Hutch. ex Gibbs, J. Linn. Soc., Bot. 42 (1914) 136. Type: Gibbs 4295 (BM, not seen), Malaysia, Sabah, between Koung and Kabayo.
Tree, 1517 m tall, with simple architecture of the main trunk and secondary branches only; twigs 78 mm in diam., glabrous, densely glaucous, bluish-green, hollow, housing ants. Bark smooth, hoop-marked, often with stipule bases persistent, bluish-green even when mature, glaucous. Stipules elliptic, 912 by 2.54.5 mm, coriaceous, spreading, not producing food-bodies, all pairs usually persistent on the shoots after leaf fall. Leaves: petioles terete, slender, 1030 cm long, shorter towards shoot apex, glabrous, glaucous, strongly double pulvinate; blades ovate, 1330 by 1220 cm, very deeply and narrowly trilobed, dissected almost to the petiole insertion or 4/57/8 of the leaf length, leaf lobes elliptic, 2.54 cm wide, ± equal in size, central lobe slightly constricted near the base and becoming spathulate, lateral lobes slightly asymmetrical and ascending and curved towards the slightly longer central lobe, 24 cm peltate, base rounded, margin entire, usually slightly revolute, with small protruding, conical nectaries, apices finely acute to acuminate, adaxial surface glabrous, abaxial surface sparsely villous with fine silvery hairs, densely glaucous; 1° venation palmate with 810 prominent veins, 2° venation strongly looping and meeting the adjacent veins near the leaf margin, 3° and 4° veins not or only weakly scalariform; young leaves dark red-brown, scattered with long silvery hairs and ferrugineus scaly hairs, producing food-bodies on lower surface. Staminate inflorescences particulate, erect, 720 by 510 cm, with minute ferrugineus hairs, 3 axis orders, main axis flattened, first pair of secondary branches ± opposite with accessory branches; bracts elliptic, c. 6 by 2.5 mm, margin entire, apex acute, covered in minute ferrugineus scaly hairs, caducous; flower clusters with c. 10 flowers, decussately arranged and crowded on ultimate inflorescence branches; bracteoles ovate, 36 by 34 mm, enclosing the flower clusters, margin finely evenly pectinate with 1016 teeth and the apical one slightly elongated, apex acute, densely covered with minute ferrugineus hairs. Staminate flowers c. 1 mm long, pedicellate; sepals fused, splitting to c. 1/4, apex densely minutely ferrugineus hairy; stamens 1; anthers 4- (rarely 3-)locular. Pistillate inflorescences narrowly paniculate, erect, 514 by 25 cm, slender, up to 2 axis orders, secondary branches ± opposite, covered in minute ferrugineus scaly hairs; bracts elliptic, c. 57 mm long, entire, covered in ferrugineus hairs, caducous. Pistillate flowers c. 23 mm long, solitary in bracteole axils; calyx urceolate, c. 1.5 mm long, minutely ferrugineus hairy, persistent; ovary 3-carpellate, 12 mm long; styles c. 1 mm long, free, spreading, persistent; stigma not dissected. Fruits subglobose, 4.56 by 56 mm, pedicellate, glaucous, bluish green, two discrete glandular patches on each carpel wall developing into slightly raised ridges, covered in bright yellow, sticky exudate; pedicel 511 mm long, minutely ferrugineus hairy, strongly glaucous. Seeds ± spheroidal or slightly flattened, c. 3 mm in diam., black, with large shallow round pits and a small centrally pointed linear scar, encased in a fleshy bright red aril.
Distribution Borneo (Sabah, Brunei, Sarawak, W, Central and E Kalimantan).
Habitat & Ecology A small-statured early successional tree very common in the lowlands of Borneo. The species is very high-light demanding with very high potential growth rates. It reaches reproductive maturity within five years. It occurs in both primary and secondary mixed dipterocarp forests. It appears to have a relatively wide tolerance of soil conditions, occurring on heavily degraded sites as well as on more fertile sites. It is uncommon in flooded or swampy areas.
Notes 1. Davies (2001) has not seen the type of M. hypoleuca var. borneensis, but the description undoubtedly refers to M. beccariana.
2. The very distinctive deeply dissected, trilobed leaves of M. beccariana, being somewhat reminiscent of cassava leaves, make this species readily identifiable. It is closely related to M. hypoleuca, under which it was previously considered a subspecies. However, the two species are often sympatric and are clearly distinct. Macaranga beccariana is a smaller tree at maturity than M. hypoleuca. In addition, M. beccariana has an extremely characteristic and unusual architecture; unless damaged the main axis produces only secondary branches, and the branch apices all come to lie at one height through elongation of the basal branches.
3. Both M. beccariana and M. hypoleuca undergo a rather protracted juvenile stage of heteroblastic leaf development, in which the leaves change slowly from entire seedling leaves with extrafloral nectaries, through partially lobed leaves, to mature lobed leaves. During this stage the species are rather difficult to distinguish. The seedling leaves of M. beccariana are diagnostic in being very dark red-brown, entire and very strongly minutely bullate, a feature not found in other Bornean Macaranga species (see M. hypoleuca below). As with M. hypoleuca this species becomes inhabited by ants at c. 10 cm tall, with the stem swelling automatically.
Macaranga belensis L.M.Perry, J. Arnold Arb. 34 (1953) 200; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 140; Whitmore, Gen. Macaranga Prodr. (2008) 88. Type: Brass 11212 (holo A); Indonesia, Papua, Lake Habbema.
Small tree to 12m, softly pilose to subvillose with hairs to 2mm long. Twigs ? Stipules 1.12.3 cm, soon caducous. Leaves: petioles 13.5 cm; blades ovate, 513 by 25.5cm, narrowing to a retuse or emarginate base with 26 glands near petiole insertion, tapering gradually to a long 12.5cm acuminate tip, drying olive above, glabrous below with pale slightly pilose nerves, yellow granular glandular on both surfaces [sic ?], with 3 nerves from petiole insertion, the laterals extending 1/3 the lamina length. Staminate inflorescences 57.5 cm spikes; peduncle 12.5 cm; flower clusters spaced; bracteoles ovate, long-acuminate, pilose outside, without patellar glands inside. Staminate flowers in clusters of 710, pilose; stamens 1124, anthers 4-locular. Pistillate parts, fruits and seeds unknown.
Distribution Indonesian New Guinea (Papua; known from type only).
Habitat & Ecology Secondary forest, common; 2200m.
Notes 1. Only known from the type, not seen by me. The somewhat inadequate (and equivocal) description is translated from Perry.
2. The absence of patellar glands from the male bracteole is unusual in this species group.
Macaranga bicolor Mόll.Arg., Linnaea 34 (1865) 199; Linnaea 34 (1866) 1010; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 319; Merr., Philipp. J. Sc. 1, Suppl. (1906) 80; Enum. Philipp. Flow. Pl. 1923) 440; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 34; Whitmore, Gen. Macaranga Prodr. (2008) 89. Type: Cuming 1301 (iso: CGE, K, L, P, W), Philippines, Luzon, Albay Prov.
[Macaranga utilis Elmer ex Merr., Enum. Philipp. Flow. Pl. (1923) 440, nomen nudum.]
Small tree to 6 m. Twigs fairly stout, 1 cm, strongly angular, finely grey-brown furfuraceous. Stipules tiny, ovate, 2 mm, apex acute, papery, densely furfuraceous, erect, soon caducous. Leaves: petioles 517 cm, slender, terete, velvety; blades ovate, 818(26) by 713(18) cm, thinly leathery, base strongly peltate, rounded, apex acute, grey below when alive; drying bicoloured: grey below, brown above or vice versa, below with sparse dark granular glands, nerves very finely furfuraceous and also sometimes with white hairs, sometimes glaucous grey below. Staminate inflorescences crowded, sometimes rather open, racemes, rusty scurfy, to 16(20) cm with axes slender; branches alternate, to 4 cm, with usually crowded small 3 mm clusters, occasionally with axillary branches, rarely with short 3rd order branches; bracts ovate, 1 mm, apex acute, furfuraceous, usually persistent; bracteoles persistent, roundish, 2 mm, apex broadly acute, margin entire to shallowly dentate, rusty scurfy on both surfaces. Staminate flowers 412 per cluster; sepals joined at base, stamens 2 or 3, anthers 4-locular. Pistillate inflorescences amongst and behind leaves, crowded, short, 4 cm, finely furfuraceous spikes, or with 5 mm branches; bracteoles and flowers not seen. Fruits solitary, often crowded, depressed globose, to 7 mm diam., (weakly) 6(8) shouldered, 3(4) locular, wall smooth, leathery, finely granular glandular except for sutures, sometimes slightly furfuraceous; pedicel to 12mm, finely furfuraceous; sepals small, persistent, separate; stigmas small, rounded, 1 mm, apically smooth. Seeds 3, roundish, pitted.
Distribution Philippines (Luzon, Mindoro, Leyte, Samar, Sibuyan, Mindanao).
Habitat & Ecology Lowlands.
Macaranga bifoveata J.J.Sm., Nova Guinea Bot. 8 (1912) 790, t.139; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 377; L.M.Perry, J. Arnold Arb. 34 (1953) 216; Whitmore, Gen. Macaranga Prodr. (2008) 89. Syntypes: Gjellerup 209 (L), Indonesia, Papua, North Coast, Salzquelle, Beguwri River; Schlechter 14113 (P), Papua New Guinea, Madang Prov., auf dem Wege von Ranu [Ramu?] zur Kόste.
Macaranga latifolia L.M.Perry, J. Arnold Arb. 34 (1953) 217.
Treelet or small tree to 10 m. Twigs slender 3 mm diam., angled and sparsely ridged near tip, pubescent. Stipules triangular, broader than long, 2 by 3 mm, apex acuminate, erect, very closely adpressed, extremely inconspicuous, patchily persistent, sparsely tomentose, leaving a scar on falling. Leaves: petioles 418 cm, slender, terete, shallowly striate, sparsely pubescent, kneed at top; blades almost as broad as long, ovate 1221 by 1023 cm, papery, base broadly truncate or very broadly shallowly cordate, at petiole minutely peltate or with 2 tiny lobes and with 2(4) large marginal sometimes depressed glands, apex broadly acute, or rounded with a 15mm acuminate tip, midbrown, nerves below variably pubescent with short stiff white often tufted spreading hairs, and variably granular glandular, lamina with diffuse golden granular glands and occasionally slightly glaucous, with 3 basal nerves. Staminate inflorescences diffuse racemes or panicles (with 4 axis orders) to 20 cm long, secondary axes well-spaced, normal to main axis, 47 cm long; peduncle 36 cm; main axis to 2 mm diam., slightly flattened at peduncle, strongly so distally, sparsely ridged, sparsely pubescent and sometimes usually sparsely granular glandular; basal secondary branches opposite, rarely a few axillary branches more distally; bracts not seen; ultimate branches slender (not thread-like); flower clusters in groups of 13 evenly spaced, almost contiguous or to 4 mm apart, 3 mm diam., 17 flowered; bracteoles trapezoid, 2 by 2 mm, spreading, persistent, smaller than cluster, granular glandular outside, thick with several small sunken patellar glands inside. Staminate flowers globose, 0.5 mm diam., shortly stalked; sepals 3, glabrous; stamens 35, anthers 4-locular. Pistillate inflorescences spikes or sparsely branched racemes to 7 cm, main axis 2 mm diam., terete, weakly striate, sparsely tomentose; peduncle 23 cm; branches alternate or basal ones opposite, 1 cm; bracteoles caducous, subrhombic-spathulate, to 4.5 mm long, puberulous outside, with several patellar glands inside. Fruits in 2 cm spaced clusters of 13, or sometimes just at apex, globose, 9mm diam., bilocular, leathery, densely uniformly covered with fine bristly 4 mm pubescent spines, tending to rub off and with a grey to green cast; pedicel stout, 6 mm, tomentose; calyx caducous; styles 23 mm, recurved, plumose. Seeds somewhat flattened, coarsely verrucose, with sarcotesta.
Distribution Throughout New Guinea.
Habitat & Ecology Usually in secondary or disturbed rain forest. Lowlands to 1500m.
Notes Distinctive in its leaf shape (especially base), stipules and fruits. The descriptions of the pistillate bracteoles are copied from Perry (1953).
Macaranga brachytricha Airy Shaw, Kew Bull. 23 (1969) 103; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)140; Gen. Macaranga Prodr. (2008) 91. Type: Hoogland 4899 (holo K; iso L), Papua New Guinea, Madang Distr., Mawau Village, Gogol Valley, 25 km inland.
Tree to 12m. Twigs stout c. 7mm diam., strongly angled, sometimes glaucous, glabrous or pubescent. Stipules elongate triangular, 2025 by 1215 mm, sides straight, apex acute, with faint parallel nerves, warm brown, papery, at first weakly scurfy, becoming glabrous, recurved, persistent. Leaves: petioles 924 cm, to 5 mm diam. near base, terete, faintly ridged, soon becoming glabrous, strongly constricted at base, weakly kneed at top; blades ovate, 1930 by 1124 cm, papery, base broadly (shallowly) rounded, 46 cm peltate, margin entire, apex 2 cm caudate, drying pale brown, above with dense small clustered hairs, below with clustered and simple hairs on all nerves and inconspicuous golden granular glands, spider's web nervation, all minor nerves scalariform, ultimate order irregularly so. Immature Staminate inflorescences narrow 9 cm panicles with 3 axis orders, at first pale furfuraceous; main branches ascending, alternate; peduncle 1 cm, slightly angular, 1.5 mm diam.; bracts variable, strap-like, oblong, to 5 by 2mm, apex acute, at first pale furfuraceous, dark brown, papery, erect, caducous; bracteoles trapezoid, 4 by 3 mm, apex acute, widest in middle, deeply narrowly regularly pectinate, both surfaces velvety, thick, erect, persistent, overlapping, longer than cluster of c. 4 flowers. Staminate flowers puberulous with c. 6 stamens, anthers 4-locular. Infructescences short axillary racemes to 5 cm long, sparsely hairy; peduncle 4 cm, angular near tip with 1-several fruit clusters on short 5 mm branches; fruits enclosed in velvety regularly deeply pectinate, ovate, 8 by 10 mm, erect papery bracts with rounded apex, with several prominent nerves. Fruits (immature?) 13 lobed, lobes globose c. 5mm across, thinly woody with sparse 3 mm spines on upper part, and wall with small warts; pedicel to 7mm, glabrous; calyx persistent; style(s) apical to 3 mm, smooth, spreading, caducous. Seeds 5 mm diam., coarsely verrucose, with sarcotesta.
Distribution East Papua New Guinea, Bismarck Archipelago (New Britain).
Habitat & Ecology Secondary forest. Lowlands.
Macaranga brevipetiolata Airy Shaw, Kew Bull. 19 (1965) 326; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 154; Gen. Macaranga Prodr. (2008) 92. Type: Kostermans 6433 (holo K; iso L), Indonesian Borneo, Loa Djanan, W of Samarinda.
Treelet or tree 210(20) m. Twigs solid, 3 mm diam., angled, striate, pubescent. Stipules needle-like, narrowly elliptic, 8 by 1 mm, including a long caudex, pubescent, stiffly erect, crowded, persistent. Leaves: petioles short, 23 cm, terete, densely pubescent, kneed at both ends; blades obovate, 818 by 47 cm, thinly leathery, base long tapering to a minutely cordate part with 2 conspicuous glands, margin sometimes slightly undulate, apex abruptly tapered to a 5 mm acute tip, often drying greyish green, below nerves densely pubescent, densely granular glandular. Staminate inflorescences pubescent spikes to 12 cm, axis terete, peduncle short, spaced flower clusters subtended by persistent bracteoles; latter elliptic, 3 by 1 mm, erect, apex acute, densely pubescent outside. Staminate flowers 34 developing in sequence, shortly pedicellate, 1.3 mm diam.; sepals pubescent; stamens c. 12, anthers 4-locular. Pistillate inflorescences as staminate, spikes to 7 cm; peduncle 34 cm. Pistillate flowers 37. Fruits solitary, subtended by one caducous variable leafy bract, ovate to elliptic, to 8 by 3 mm, pubescent, spreading; fruits strongly dicoccous, each lobe round to 5 mm diam., densely pubescent with 3 mm flattened slightly pubescent linear spines; pedicel 3 mm; calyx small, persistent, pubescent; styles thread-like, 2 cm, soon caducous. Seeds smooth, without sarcotesta.
Distribution Throughout Borneo.
Habitat & Ecology Primary forest understorey species, also present in selectively logged forest. On sandy loam, sandy clay or dry sandstone soils (common at Lambir, Sarawak; S.J.Davies, pers. comm.)
Macaranga brunneo-floccosa Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv.add.VI (1919) 28; L.M.Perry, J. Arnold Arb. 34 (1953) 256; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 140; Gen. Macaranga Prodr. (2008) 92. Syntypes: Ledermann 12809, 13035a (not seen), Papua New Guinea, Sepik Region, Felsspitze.
Macaranga brunneo-floccosa Pax & K.Hoffm. var. calvescens Pax & K.Hoffm in Engl., Pflanzenr. IV.147.xiv.add.VI (1919) 28
Trees, 1020(33)m tall, or rarely a 3 m shrub. Twigs slender, 23 mm diam., terete, striate, at first rufous floccose. Stipules elliptic, to 5 by 2 mm, apex acute, warm brown, papery, sparsely tomentose, erect, soon caducous. Leaves: petioles 14 cm, slender, 1 mm diam., terete, at first rufous floccose, kneed at both ends; blades ovate to ovate-elliptic, 49(13) by 2.55.5(7)cm, leathery, base rounded, drying pale golden brown, margin entire, apex abruptly acuminate, at first rufous floccose below and near petiole insertion above becoming glabrous, without granular glands, secondary nerves only (3)4(5) pairs, distant, curved, looping and joining near margin, basal pair submarginal, emerging from petiole insertion, sometimes somewhat concave, or sometimes coarsely bullate, reticulations slightly raised on upper surface. Staminate inflorescences dense spikes or racemes to 4 cm or 7 by 4 cm panicles with 34 axis orders, usually rufous floccose, branches divaricate or ascending; peduncle 210 mm; main axis 1 mm diam., terete;, lower branches rarely opposite; flower clusters irregularly spaced, contiguous to 3 mm; bracteoles broadly ovate, 1.2 by 1.5 mm, apex blunt, cucullate, papery, spreading, pubescent outside, smaller than cluster. Staminate flowers in clusters of 6, sessile; calyx pubescent; stamens 6, anthers 4-locular. Infructescences 4 cm, rufous floccose spikes with 13 fruits. Fruits 8 mm diam., shallowly bilobed, wall thick, toughly fleshy, blackish with evanescent rufous tomentum; pedicel stout, 10 mm; calyx persistent; styles 8 mm, twin, erect apical, stout, persistent, papillose. Seeds subglobose, 3 by 4.5 mm, coarsely rugose, with sarcotesta.
Distribution Scattered across New Guinea.
Habitat & Ecology Primary montane forests, 8002100m.
Notes 1. Highly distinctive in its leaves with few secondary nerves and absence of granular glands. Very variable in leaf size and male inflorescence construction. Despite this polymorphism I believe this is all one species, though when more material is available vicariant subspecies might be discernable.
2. I confirm Perry's observation (1953) that the inside ovary wall of Brass & Versteegh 13163 is densely pubescent, a condition also seen in M. carrii and M. sterrophylla; van Royen & Sleumer 8095 is glabrous as normal.
Macaranga caladiifolia Becc., Malesia 2 (1884) 46; S.J.Davies, Harvard Pap. Bot. 6 (2001) 393. Macaranga caladiifolia Becc. var. pilosula Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 384, nom. inval. Type: Beccari PB 920 (holotype: Fl, not seen), Malaysia, Sarawak.
Macarunga puncticulata auct. non Gage: Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 383; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 150.
Small unbranched or once-branched treelet, 13 m tall; twigs 612 mm in diam., glabrous, prominently swollen and constricted at some but not all nodes, hollow, housing ants. Stipules narrowly ovate, 813 by 35 mm, apex acuminate, glabrous, the pair not completely encircling the twig, coriaceous, spreading, producing food-bodies on the abaxial surface, up to 5 pairs present on the shoots. Leaves: petioles terete, slender, 822 cm long, glabrous; blades narrowly ovate, 1528 by 714 cm, entire or sometimes with small lateral cusps, 46 cm peltate, base rounded to almost truncate, margin entire to quite coarsely dentate, with prominent upward-facing flattened nectaries particularly towards the leaf base. apices finely acuminate, adaxial surface glabrous and glossy, abaxial surface glabrous or with minute hairs scattered along veins, not glaucous; 1° venation palmate with 57 prominent veins, 2° veins looping strongly near the leaf margins, 3° venation very weakly scalariform; young leaves reddish-brown, scattered with minute ferrugineus hairs on upper surface, lower surface densely glandular punctate. Staminate inflorescences narrowly paniculate, erect, to c. 15 cm long, glabrous, red, 2 axis orders, the first pair of opposite branches arising at the apex of a long and slender main axis; bracts ovate, 58 by 34 mm, persistent, margin entire, apex acute, glabrous; flower clusters with 15 flowers, spirally arranged and crowded at the apex of the inflorescence; bracteoles ovate, 35 by 24 mm, delicate and thin, enclosing flower clusters, margin ± entire to slightly wavy, apex acute to acuminate, glabrous or sometimes with a few scattered ferrugineus hairs at the base. Staminate flowers 12 mm long, pedicellate; sepals free, very thin, glabrous; stamens (3)45; anthers 4-locular. Pistillate inflorescences erect, unbranched, with the flowers clustered at the distal end of the rhachis, 25 by c. 1 cm, stout, glabrous; bracts caducous. Pistillate flowers c. 1.5 mm long, solitary in bract axils; calyx urceolate, c. 1 mm long, scattered with fine red-brown hairs, persistent, splitting irregularly as ovary expands; ovary 3-carpellate, c. 1 mm long; styles c. 23 mm long, fused at base, free and spreading from c. 1/3 of the length, persisting to form a prominent 24 mm long crown at the fruit apex; stigma not dissected. Fruits subglobose, c. 8 by 10 mm, bright red, shortly pedicellate, two discrete glandular patches on each carpel wall developing into slender ridges c. 2 mm high, covered in pink, sticky exudate; pedicel 45 mm long, covered in fine red-brown hairs. Seeds subtriangular-ovoid, 5 mm in diam., brown-black, with shallow narrow grooves and a small cruciform scar, encased in a fleshy brightly orange-red aril.
Distribution Borneo (West Kalimantan and Sarawak).
Habitat & Ecology An understorey treelet in heath (kerangas) forest and slightly wetter (kerapah) forest over nutrient-poor acidic sandy soils. Collected in kerangas forest from sea level in Kubah National Park in west Sarawak to 450 m above sea level in kerangas forest on the summit of the hills in Lambir Hills National Park in northern Sarawak. The species is very shade tolerant, and has not been seen in degraded or logged forests as is common with many of its close relatives.
Notes 1. Davies (2001) has not seen the type specimen, but the illustration with the original description is excellent and perfectly diagnostic.
2. The ecology of M. caladiifolia is highly distinctive and unlike most other species of western Malesian Macaranga. This species appears to be closely related to M. kingii, M. lamellata and M. umbrosa with which it shares two small glandular patches on the wall of each carpel which develop into prominent ridges, and few flowers in each flower cluster. With the latter two it shares entire leaves. Macaranga caladiifolia differs from these species in having unevenly swollen stems (not all internodes are hollow), three carpellate fruits, and thin spreading stipules, as well as a distinctive ecological habit. Only M. havilandii, among Bornean Macaranga, also reproduces at around 12 m in height, but M. havilandii has broadly ovate recurved stipules, unistaminate flowers, and 45 carpellate fruits.
3. Whitmore (1975) synonymised M. puncticulata Gage with M. caladiifolia, based on inadequate material and limited field observations of these species. Macaranga puncticulata differs in many ways from M. caladiifolia; it grows to be a tree of 1520 m height and is largely restricted to peat swamp forests, furthermore it has globose evenly glandular 34-carpellate fruits, and pubescent inflorescences.
Macaranga calcicola Airy Shaw, Kew Bull. 25 (1971) 536; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 146; S.J.Davies, Harvard Pap. Bot. 6 (2001) 394. Type: Jacobs 5480 (holotype: K; isotypes: L, SAR), Malaysia, Sarawak, Ban, Sebuaran.
An elegant small tree of 410 m tall with glossy green leaves; twigs 47 mm in diam., often drying whitish, covered in long erect silvery hairs, hollow, housing ants. Bark smooth, greyish brown. Stipules ovate, 68 by 79 mm, scattered with short ferrugineus hairs at the base of the adaxial surface, the pair completely encircling the twig, succulent, reddish- to purplish brown when fresh, recurved, producing food-bodies on the abaxial surface, up to 4 pairs present on the shoots. Leaves: petioles terete, slender, 820 cm long, finely pubescent particularly towards the base and apex; blades, very broadly triangular-ovate, 1224 by 820 cm, tricusped to shallowly trilobed, dissected to 1/5 of the leaf length, central lobe triangular 615 cm wide at the base and narrowing gradually towards the apex, lateral lobes c. 13 by 13 cm at the base shortly acute and spreading, 1.54 cm peltate, bright green and shiny when fresh, base very broadly rounded, margin entire, apices shortly acuminate, adaxial surface glabrous, abaxial surface with dense short curved silvery hairs on the veins; 1° venation palmate with 78 prominent veins, 2° venation scalariform with 911 pairs arising from the midrib, looping upwards near the leaf margins and mostly terminating in minute protruding, flattened and usually downward-facing nectaries; young leaves light brown, shiny, finely pubescent on abaxial veins and finely ferrugineus pubescent when very young. Staminate inflorescences short panicles with few branches, erect, 515 by 38 cm, basal unbranched rhachis very short and only 0.53.5 cm long, basal and main axes densely covered in c. 1 mm long erect sharp-pointed hyaline hairs, distal branches covered in minute soft silvery hairs, 23 axis orders, main axis flattened, first pair of secondary branches ±opposite with or without accessory branches; bracts elliptic, 57 by c. 3 mm, margin entire, apex acute, finely pubescent particularly towards the base and on the margins, caducous; flower clusters dense with 2050 flowers, spirally arranged and clustered towards the ends of ultimate inflorescence branches; bracteoles deltate, 36 by 23 mm, margin entire, often with one or two prominent conical nectaries on the margin adjacent to and/or on the apex, apex narrowing suddenly to a slender acuminate point, only partially enclosing the flower clusters, scattered on both surfaces with minute ferrugineus hairs particularly dense towards the base of the adaxial surface. Staminate flowers c. 1 mm long, sessile; sepals fused, splitting irregularly to c. 1/5, apex densely covered in minute ferrugineus hairs; stamens 1; anthers 3-locular. Pistillate inflorescences short, stout, erect, 1.56 by c. 1.53.5 cm, usually with one pair of short opposite branches only, appearing trifid, bracts caducous, not seen. Pistillate flowers 57 by 2.53.0 mm, solitary in bract axils; calyx urceolate, apex truncate, c. 2 mm long, glabrous, persistent, splitting irregularly as the ovary expands; ovary 4- (rarely 5)-carpellate, globose, 34 mm in diam.; styles 34 mm long, rigidly erect, fused for most of the length, free and spreading from c. 4/5 of the length, persisting to form a prominent 34 mm long crown at fruit apex; stigma smooth. Fruits subglobose, 67 by c. 710 mm, sessile, one very small discrete glandular patch on each carpel wall developing into a slightly raised swelling <1 mm long, covered in light yellowish green, sticky exudate. Seeds subtriangular-ovoid, c. 4 mm in diam., black, with shallow coarse grooves and a small cruciform scar, encased in a fleshy aril (colour not seen.).
Distribution Borneo (Sarawak).
Habitat & Ecology A small early successional tree endemic to the limestone formation of Bau in western Sarawak. It is only known from low altitudes. It is a relatively small tree and is common at the base, and often grows along the sides, of steep rocky karst limestone.
Note This species is a member of the core group of section Pachystemon with unistaminate flowers and 3-locular anthers. The small glossy tricusped leaves, the gland-tipped staminate bracteoles, and the glandular fruit walls lacking prominent horn-like processes distinguish this species from its close relatives. It has several features suggesting a close relationship with M. petanostyla including glossy leaves, and staminate bracteoles that do not fully enclose the flower clusters (see further discussion under that species).
Macaranga carrii L.M.Perry, J. Arnold Arb. 34 (1953) 235; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 141; Gen. Macaranga Prodr. (2008) 97. Type: Carr 15288 (holo A; iso BM, K, L, SING), Papua New Guinea, Northern Prov., Mt. Ganeve.
var. carrii
var. carrii: Whitmore, Gen. Macaranga Prodr. (2008) 97. Type: as the species.
Tree 624m. Twigs stout, 46 mm diam., squarish, densely orange-brown tomentose at first. Stipules ovate, 18(25) by 10(18) mm, apex rounded or acute, blackish, papery, with orange-brown tomentum at least along centre, erect. Leaves: petioles 38 cm, diam. 1.54 mm, angled, striate, variously orange-brown tomentose, kneed at top; blades ovate to deltoid, size variable, 716 by 514 cm, leathery, base broadly rounded to truncate, sometimes shallowly broadly cordate, margin with distant small glandular teeth, apex a broad 15 mm caudex, 6 mm broad at its base, drying dark grey-brown above and glabrous except near base of midrib, at first densely orange-brown tomentose below, becoming glabrous to expose black granular glands, very strongly and finely doubly bullate between both reticulations and the scalariform tertiary nerves, base with 3 major and several minor nerves arising from petiole, below all nerves strongly raised. Staminate inflorescences narrow intertwined orange-brown pubescent 10 cm panicles; branches short to 5 cm, alternate, more or less ascending, sinuous, distally thread-like and zigzag and bearing a few 1 cm tertiary branches, some branches arising from a flower cluster; peduncle 2 cm, main axis distally slender, at base stout, 2 mm diam., terete, densely pubescent; bracts subpersistent, variable, 7 by 24 mm, oblong or widening towards rounded tip (or with 3 narrow lobes), patchily tomentose without, blackish, papery, flower clusters to 5 by 3 mm, evenly spaced, sometimes in opposite pairs, 8 mm apart or towards branch tips contiguous; bracteoles broadly elliptic, 5 mm, 1.5 mm wide near base and long tapering acuminate, base rounded, margin irregularly serrate, hairy, outside with sparse long hairs, papery, spreading slightly beyond cluster, persistent. Staminate flowers 6 per cluster, large, 2 mm diam., calyx with long sparse hairs and granular glands, stamens 7, 4-locular, sessile. Infructescences open, sparsely branched 15 cm racemes, branches sinuous 25 cm, peduncle 24 cm, with stipule-like caducous bracts. Fruits solitary, 3 mm apart, as 1 cm diam., leathery, softly spiny woolly balls, bilocular, the inner ovary wall hirsute; pedicel stout, 2 mm; calyx inconspicuous, persistent; styles 2, recurved, 8 mm, stout (1 mm), papillose, arising from a 1 mm fused base. Seeds (young) shallowly verrucose, with sarcotesta.
Distribution Papua New Guinea (Western Highlands, Southern Highlands, Central Prov.).
Habitat & Ecology Forest edges and regrowth. Mountains c.2400m.
Notes 1. The staminate inflorescence bracts are variable, but only Perry (1953) has seen trilobed ones. The staminate sepals I examined are not trilobed pace Perry. Inner fruit wall hairy also seen in M. brunneo-floccosa and M. sterrophylla.
2. A striking species of montane forests with the leaves ovate, coriaceous, coarsely rufous tomentose and prominently bullate (except var. myolensis). Known from five partly sympatric varieties which are only narrowly distinct, by the characters shown in the key, and when more collections have been made may perhaps merge. Macaranga clemensiae (see below) is close, but does not differ between its several localities. Macaranga tanarius is one of a group of closely allied species, which also form a knot of closely related taxa.
var. laevis Whitmore
var. laevis Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 141; Gen. Macaranga Prodr. (2008) 98. Type: Gillison NGF 25144 (holo K; iso L, SING), Papua New Guinea, Soutern Highlands Dist., Tari Gap, Ibiwari.
Differs in Leaves with base 410 mm peltate, apex acute only rarely caudate, tomentum below sometimes dark chocolate brown. Staminate inflorescences with big 510 mm diam. globose clusters; bracteoles big, broadly ovate, 5 by 6 mm, apex rounded, fleshy, hirsute outside, glabrous inside, spreading, not extending beyond cluster. Staminate flowers with thick filaments, tapering to a short stout pedicel. Fruits tomentose but not spiny.
Distribution Papua New Guinea (part of the central mountains: Western, Eastern and Southern Highlands, Star Mountains)
Habitat & Ecology Primary, disturbed and secondary montane forests, 22502900 m.
Note Described on its smooth fruits, but the big male clusters and bracteoles are also very distinctive.
var. leonardii (L.M.Perry) Whitmore
var. leonardii (L.M.Perry) Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 141; Gen. Macaranga Prodr. (2008) 98. Macaranga leonardii L.M.Perry, J. Arnold Arb. 34 (1953) 234. Type: Brass 10969 (holo A), Indonesia, Papua, Lake Habbema.
Differs in Tree 2028m. Stipules carinate especially towards tip. Leaves: blades with base (4)710(15) mm peltate, very stiff. Staminate inflorescences with bracteoles broadly ovate, 2.5 mm long. Staminate flowers very big, 5 mm long after anthesis. Pistillate flowers with unilocular ovary.
Distribution Indonesian New Guinea (northern and southern sides of Doormantop range: Lake Habbema, Idenburg river)
Habitat & Ecology Primary and secondary mountain forests, 22002880m.
Note Only known from 3 collections, none seen by me.
var. myolensis Whitmore
var. myolensis Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 142; Gen. Macaranga Prodr. (2008) 99. Type: Croft & Lelean LAE 60529 (holo K; iso L), Papua New Guinea, Central Prov., Lake Myola.
Differs in Leaves not at all bullate, base 1 cm peltate, secondaries only slightly raised below and tertiaries less so. Staminate inflorescences densely congested, peduncle and lower main axis very stout, 4 mm diam., clusters round, 5 mm diam.; bracteoles oblong, 4 by 2.5 mm, apex acute, thick, fleshy. Staminate flowers c. 20 per cluster, 1.5 mm diam., shortly pedicellate.
Distribution Papua New Guinea (Central Distr.).
Habitat & Ecology Forest edge. Mountains, 1900m.
Note Known from a single male collection, sympatric with var. carrii.
var. womersleyi (L.M.Perry) Whitmore
var. womersleyi (L.M.Perry) Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 142; Gen. Macaranga Prodr. (2008) 99. Macaranga womersleyi L.M.Perry, J. Arnold Arb. 34 (1953) 232. Type: Womersley NGF 4475 (holo A; iso K, L, SING), Papua New Guinea, Western Highlands Dist., near Nondugl.
Differs in Stipules elliptic oblong, smaller, 8 mm, apex somewhat obtuse; blades with base minutely 2 mm peltate and usually a few small glands near petiole insertion. Staminate inflorescences with 5 mm diam. clusters of 20 flowers; bracteoles ovate, 1.5 by 2 mm, apex blunt, fleshy, smaller than cluster. Staminate flowers small, 1 mm diam.
Distribution Papua New Guinea (Western Highlands Prov.).
Habitat & Ecology Mountain forest, 1950 m.
Note Still known only from a single male collection. Closest to var. carrii, and sympatric with var. laevis.
Macaranga caudata Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv add. vi (1919) 30; L.M.Perry, J. Arnold Arb. 34 (1953) 213; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)142; Gen. Macaranga Prodr. (2008) 100. Type: Ledermann 6616 (iso K), Papua New Guinea, Sepik Region, Malu.
Shrubs, 4.5 m or small trees to 10 m. Twigs slender, angular, to 4 mm diam., tomentose to subglabrous with harshly irritant hairs, occasionally glaucous, with stipule scars. Stipules united, apparently single, surrounding the apical bud, to 2 cm long, tapering from 59 mm base, golden-tomentose, papery, erect, subpersistent, sometimes splitting at tip. Leaves: petioles 615 cm, slende, 2 mm diam., terete, tomentose, weakly kneed at top; blades orbicular deltoid, 1223 by 1020 cm, papery, base broadly cordate to truncate, cordulate at petiole insertion, with few-numerous small glands, margins with small prominent glandular teeth, apex 27 cm caudate, drying fawn, both surfaces feeling velvety, above with long single hairs arising from small papillae, below nerves with spreading mostly single, a few tufted, white hairs, granular glands regular close dark, or patchy same colour as lamina, secondary nerves pinnate, basal pair from petiole insertion. Staminate inflorescences slender narrow racemes or panicles to 13 by 5cm with 3 orders of branching, main axis flattened, 1.5 mm across, pubescent; peduncle 5 cm; secondary axes divaricate, opposite to 3 cm long, thread-like, zigzag between tiny evenly spaced flower clusters, tertiary axes absent or to 5 mm long; bracteoles with 0.5 mm broad stalk subtending flowers, widening to 1.5 mm diam., roundish, apex blunt to broadly acute, spreading, fleshy, nerved, margin crenate, head bearing several sunken glands, glabrous. Staminate flowers in clusters of 14, tiny, subsessile, stamens 3, anthers 4-locular. Pistillate inflorescences long spikes to 30 cm, axis somewhat flattened, 2 mm across, furfuraceous or pubescent; peduncle 26 cm, a few near-apical clusters of a few flowers. Very young Fruits globose. densely golden pubescent with 0.5 mm close spines; pedicel >1mm long; calyx persistent; styles stout, apical, caducous. Seeds unknown.
Distribution New Guinea.
Habitat & Ecology Primary or disturbed forest; lowlands to 1000 m.
Note The leaf hairs above are reminiscent of M. hispida.
Macaranga celebica Koord., Meded.'s Lands Plantent. 19 (1898) 626; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii(1914) 392; Whitmore, Gen. Macaranga Prodr. (2008) 101. Syntypes: Koorders 16914β (BO ,L); 16978β (BO, L), Sulawesi.
Small trees to 15m. Twigs round or angled, glabrous, sometimes blackish sometimes glaucous, fairly stout, 8 mm diam. Stipules ovate, 8 by 3 mm, apex acute, dark brown, at first rufous pubescent, becoming glabrous, papery, erect, soon caducous. Leaves: petioles 1015 cm, fairly slender, round, glabrous, kneed at top; blades ovate elliptic, broadest near base, 1624 by 511cm, thinly leathery, base broadly rounded minutely peltate, drying grey above, margin entire, apex acuminate, glabrous, grey green below, closely golden granular glandular below, secondary nerves few, 810 pairs, distant (c. 2 cm), conspicuous, basal pair close and parallel to margin, tertiaries scalariform, they and sometimes minor reticulations clearly visible on both surfaces, with lines of several roundish glands on each side of petiole insertion. Staminate inflorescences with 3 round, pistillate ones with 2 flattened axes orders, to 10 cm long, with lower branches spreading, slender but not filiform; basal peduncle to 2 cm, weakly furfurescent; pistillate one with minute, entire persistent bracts; staminate one with persistent bracteoles, ovate, 2 by 2 mm, apex rounded, very weakly dentate, thick, furfuraceous, longer than the crowded clusters of c. 5 flowers. Staminate flowers shortly pedicellate; stamens 3, anthers 4-locuar. Young Fruits unilocular, round, with 3 mm furfuraceous pedicel, subtended by persistent calyx, stigma apical. Seeds unknown.
Distribution Sulawesi (Minahassa, Malili, Gunung Sadaunta), Maluku (Buru).
Habitat & Ecology Lowlands to 1200m.
Note The leaves are very distinctive, especially in their conspicuous distant secondary nerves.
Macaranga chlorolepis Airy Shaw, Kew Bull. 25 (1971) 530; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 142; Gen. Macaranga Prodr. (2008) 101. Type: Schodde 2849 (holo K; iso L), Papua New Guinea, Central Prov., Sirinumu, 7 miles S of Sogeri.
Trees 3.620 m tall. Twigs slender, 3 mm diam., terete, at first densely covered with tiny mustard yellowish lepidote scales, becoming glabrous. Stipules not seen. Leaves: petioles 2.54 cm, slender, 1.5 mm diam., terete, very finely scurfy lepidote, kneed at apex; blades ovate-elliptic, 1019 by 4.56cm, papery, base narrowly cuneate sometimes minutely cordulate, margin entire or weakly sinuate, apex acuminate, upper surface with several small glands, ochreous-grey below greyish above, below densely covered with minute yellowish or whitish lepidote scales, above with a few such scales at first. Staminate inflorescences single or to 3 per axil, short 3 cm catkins, completely covered by the flower clusters; bracteoles truncate saccate, 2 by 2 mm, base gibbous, entire, apex acute, glabrous within yellow lepidote hairs outside, erect, persistent, smaller than flower cluster. Staminate flowers to 6 per cluster; pedicel 1 mm; sepals 3, ovate, 2 by 1 mm, apex obtuse, yellow lepidote hairs outside; stamens 1722, anthers 4-locular. Pistillate inflorescences 24 cm spikes, with 23 apical flowers; bracts narrowly elliptic, to 4 by 1.5 mm, apex obtuse, shortly stalked, lepidote. Pistillate flowers: pedicel 1 mm; sepals as staminate ones; ovary bilocular with 2 thick styles 3 mm long with recurved tip. Fruits strongly dicoccous, each half globose, 5 mm diam., smooth, densely minutely yellowish lepidote, wall thick woody;, shortly pedicellate. Seeds round, 4 mm diam., smooth, completely enclosed in a thin sarcotesta.
Distribution East New Guinea, Bismarck Archipelago (New Britain, New Ireland).
Habitat & Ecology Primary forest 30600m.
Notes 1. The only other Macaranga with lepidote scales is M. subdentata of north Queensland. Parts of the description above are based on Airy Shaw.
2. The scales appear to be flattened granular glands.
Macaranga choiseuliana Airy Shaw, Kew Bull. 32 (1978) 412; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)1368 passim; Gen. Macaranga Prodr. (2008) 102. Type: Whitmore's Collectors BSIP 5677 (holo L; iso K), Solomon Islands, Choiseul, banks of Vach River.
Trees, sometimes reaching 30 m tall, 60 cm diam. Twigs very stout, to 2 cm diam., irregularly roundish, with big conspicuous leaf and stipule scars, velvety tawny tomentose with white crisped simple and tufted hairs. Stipules big, elliptic, to 13 by 2.5 cm long, tapering to an acuminate apex, tawny brown, papery, somewhat crumpled, velvety with dense tomentum of simple and tufted hairs, crowded, persistent, forming an erect bundle protruding beyond apical bud. Leaves: petioles massive, 45 cm long, stout, 8 mm diam., flattened above, tomentum as stipules, kneed at top, thin, base often contracted; blades big, ovate, 3050 by 2040 cm, base rounded, to 12 cm peltate, margin repand, usually with close small protruding glandular teeth, apex acuminate to very shortly caudate, drying khaki above, tawny below, above main nerves tomentose, below velvety, all nerves with white simple or tufted hairs and lamina black granular glandular, with spider's web nervation, tertiaries raised below, scalariform. Staminate inflorescences dense pyramidal 7 cm panicles with 3 axis orders; peduncle 1 cm, emergent from a tuft of roundish 8 mm papery bracts, flattened 1.5 mm diam., tomentose with spreading white hairs; secondary axes alternate, some with axillary branches, to 3 cm, crowded and intermingling, golden pubescent; flower clusters tiny, 1 mm diam., almost touching; bracteoles tiny, rounded ovate, 1 by 0.5 mm, apex blunt, thick, spreading, adaxial surface glabrous, abaxial surface fufurfaceous, persistent, smaller than cluster. Staminate flowers 7 per cluster tiny, 0.5 mm, sessile; sepals pubescent; stamens 12, anthers 4-locular. Pistillate inflorescences small dense panicles, 815 cm long, with 3 tawny velvety axis orders, branches crowded, steeply ascending, secondaries 6 cm long, sinuous, slender, occasionally with axillary branches, tertiaries to 5 mm; peduncle short, 12 cm, stout, squarish to flattish, 3 mm diam., with several soon caducous tawny velvety rounded papery bracts to 8 by 5 mm; main axis and lower secondaries stout, secondaries subtended by 2 by 3 mm ovate, rounded, velvety persistent small spreading bracts, flower clusters evenly 25 mm spaced. Pistillate flowers solitary, rarely twinned; bracteoles minute, ovate, tomentellose; calyx cupuliform, tomentellose. Fruits strongly bilobed, slightly flattened, 5 by 8 mm, thinly woody, smooth, dark brown with evanescent tawny furfurescent scurf; pedicel stout 3 mm, tawny scurfy; calyx small, persistent; styles 2, apical 1 mm recurved, persistent Seeds globos, 4 mm diam., coarsely regularly verrucose, with sarcotesta.
Distribution Papua New Guinea (Bougainville), Solomon Islands (Choiseul).
Habitat & Ecology Well drained primary lowland rain forest.
Notes 1. The staminate bracteoles are so tiny as to be little more than outgrowths of the axis; they are glandless pace Whitmore (1980). Airy Shaw described the fruits as sometimes partly minutely warty, but this feature is so minimal that I omit it.
2. Sympatric in Choiseul with M. whitmorei which, however, is almost glabrous and has long 5 mm styles on the fruits.
Macaranga chrysotricha Lauterb.& K.Schum. in K.Schum. & Lauterb., Fl. Schutzgeb. Sόdsee (1900) 399; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 378; L.M.Perry, J. Arnold Arb. 34 (1953) 226; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 143; Gen. Macaranga Prodr. (2008) 103. Type: Bamler II 22 (holo B, lost), Papua New Guinea, Morobe Prov., Sattelberg.
Macaranga chrysotricha var. glaucescens Mansfeld, J. Arnold Arbor.10 (1919) 78, 233; L.M.Perry, J. Arnold Arb. 34 (1953) 227.
Small trees to 8 m, invested with long golden irritant hairs. Twigs angular, to 7 mm diam., with stipule scars, occasionally glaucous. Stipules paired, ovate oblong, to 3.5 by 2 cm, apex acuminate, chestnut brown, outside densely golden tomentose, occasionally glaucous, inside glabrous, erect, papery, subpersistent. Leaves: petioles 510 cm, terete, 3 mm diam., variously golden tomentose, sometimes weakly kneed at top; blades deltoid, 1317(36) by 914(30) cm, leathery, base very variable, truncate, shallowly broadly cordate, deeply cordate, or very deeply cordate with a narrow 2 cm broad, 7 cm deep sinus, occasionally 5 mm peltate, with several large sometimes inconspicuous glands near petiole insertion, margin often coarsely repand, usually with regular, distant glandular teeth, apex acute, occasionally 2 cm caudate, drying dark brown above, paler below but suffused by the golden tomentum, usually finely double bullate above, both surfaces golden tomentose, becoming confined to main nerves above, hairs mostly solitary to mostly tufted, below all nerves raised and tomentose, below black granular glandular or glandless, with 24 secondary nerves from petiole insertion. Staminate inflorescences long-stalked panicles to 22 cm with 3 axis orders, main axis angular to flattened, 2 mm diam., tomentose, peduncle to 15 cm; lowest branches opposite, divaricate, to 3 cm, themselves with opposite 2 cm thread-like branches with axillary branches, bearing 3 mm spaced flower clusters; bracteoles transversely ovate, 1 by 2 mm, cucullate, fleshy, hispid outside, glabrous inside, enclosing cluster, bearing centrally an oblong elliptic, 1.5 by 1.2 mm, acute, fleshy recurved tip, bearing adaxially 2 rows of 6 patellar glands and abaxially granular glandular. Staminate flowers 6 per cluster, 0.5 mm diam.; pedicel 0.4 mm, glabrous; sepals free; stamens 3, anthers 4-locular. Infructescences long spikes to 30 cm, with a few solitary fruits crowded near apex, axis terete, 3 mm diam., golden tomentose. Fruits (immature) globose, 8 mm diam., densely golden tomentose, granular glandular; pedicel stout, 3 mm; calyx persistent; styles apical, 2 mm, recurved, plumose.Seeds unknown.
Distribution Papua New Guinea.
Habitat & Ecology Disturbed or secondary, usually montane forest (450)2500m.
Notes 1. A variable species, especially in the leaves. The var. glaucescens comes within the overall range of variation.
2. Despite its twin stipules, this species clearly belongs in the Longistipulata group. The staminate bracteoles have the distinctive gland-bearing apex, but the flowers are a more normal size, not the mere flecks common in this species group. Instead of fine sinuous spines the fruits bear a golden tomentum, below which Perry (1953) claims to have seen granular glands.
Macaranga clavata Warb., Bot. Jahrb. 13 (1891 349; K.Schum. & Lauterb., Fl. Schutzgeb. Sόdsee (1900) 396; Airy Shaw, Kew Bull. 23 (1969) 99 (descr. ampl); Kew Bull. 37 (1982) 26; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 143, Pl.6 Fig.2; Gen. Macaranga Prodr. (2008) 104. Type: Warburg s.n., 1889 (not seen), Papua New Guinea, Morobe Prov., Finschhafen. Perry (1953: 240) cited the pistillate collection Warburg 20511 in A as a possible isotype of this species.
Macaranga tanarius auct. non (L.) Mόll.Arg.: J.J.Sm., Nova Guinea 8 (1912) 791; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 353, pro parte; L.M.Perry, J. Arnold Arb. 34 (1953) 240.
Small spreading trees to 10(18) m, with shortly softly tomentose and often viscid twigs, petioles and inflorescences. Twigs rather slender, 3 mm diam., angled, often with short 12 cm internodes and somewhat zigzag. Stipules broader or as broad as long, broadly triangular, to 8 by 10 mm, base truncate, often broadly auriculate sometimes subsagittate, apex acute, dark brown, papery, somewhat tomentose especially on midline, erect to spreading, persistent. Leaves: petioles to 20 cm, slender, 2 mm diam., terete, finely ridged, sometimes drying with constricted base; blades ovate, 827 by 624 cm, papery, base broadly rounded, 27 cm peltate, margin entire, apex 1.5 cm caudate, drying khaki, glabrous and shiny above, below with variously dense short spreading hairs on nerves, granular glands black (less often golden), small, inconspicuous, sometimes sparse and irregular, with spider's web nervation, tertiaries scalariform, quaternaries subscalariform to reticulate. Staminate inflorescences dense, sometimes intermingled pubescent panicles, 11 by 69 cm; main axis bearing 2 cm spaced fascicles of up to 5, 16 cm long, variously divaricate branches; peduncle 13 cm, flattened, 1.5 mm across; branches slender to thread-like, some themselves bearing fascicles of short branches; bracts triangular, to 2 mm long, pubescent, soon caducous; flower clusters irregularly spaced tending to be crowded towards branch tips; bracteoles very broadly ovate, 35 mm long and almost as broad, apex acute, strongly cucullate, densely pubescent, pectinate, persistent, longer than flower clusters of c. 10, tiny, 1 mm diam. Staminate flowers glabrous; sepals 3; stamens 6, anthers 4-locular. Infructescences spikes to 6 cm with solitary or paired fruits near the tip, axis flattened, to 2 mm across, pubescent. Fruits 12(3) lobed, lobes round, 5 mm across, thinly woody, bearing 24 mm long blackish spreading rigid viscid conspicuously clavate hairs, swollen towards apex; pedicel slender length variable to 3 cm; styles subulate, 23 mm, caducous. Seeds round, 56 mm across, coarsely regularly verrucose, with sarcotesta.
Distribution Maluku (Kai Islands), New Guinea (incl. Bismarck Archipelago).
Habitat & Ecology Secondary forests, lowlands, to 900 m.
Notes The clavate fruit processes, reminiscent of Drosera leaf hairs, are highly distinctive, as is the bizarre fascicled branching of the male inflorescences. Macaranga herculis and M. tentaculata (see below) have similar but more tentacle-like fruit processes, long stipules and the leaves densely velvety below.
2. Sayers NGF 21811 is a robust, luxuriant form with 25 by 8 cm staminate panicles and stout 1 cm diam. twigs.
Macaranga clemensiae L.M.Perry, J. Arnold Arb. 34 (1953) 232; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 143, Pl.6, Fig.3; Gen. Macaranga Prodr. (2008) 105, Fig. 5. Type: Clemens 7902 (holo A; iso K), Papua New Guinea, Morobe Prov., Samanzing.
Trees 1115 m. Twigs stout, 34 mm diam., angled, densely tomentose or pubescent. Stipules elliptic to ovate, 715 by 35 mm, apex acute, thick, densely dark brown pubescent, soon caducous. Leaves: petioles 46 cm, stout, 2 mm diam., terete, densely pubescent, kneed at both ends; blades ovate, 915 by 48 cm, thickly leathery, base broadly rounded, 25 mm peltate or not peltate, margin recurved, apex 2 cm caudate, basal nerves palmate, drying mid brown, paler above, conspicuously doubly bullate - both between secondaries and reticulations, densely to sparsely woolly tomentose below and black granular glandular, secondaries below massive, almost equalling midrib, only 47 pairs, strongly ascending, basal pair at petiole insertion and running up at least half of lamina. Staminate inflorescences pubescent or tomentose dense pyramidal panicles to 10 by 6 cm, with 3 axis orders, branches ascending to spreading; peduncle 12 cm; main axis angular, 1 mm diam.; secondaries alternate to 3 cm, sinuous; tertiaries to 1 cm, thread-like; bracts tiny, triangular, 0.8 mm, apex blunt, tomentose, erect, persistent; flower clusters close to contiguous; bracteoles obtriangular, 1 by 2 mm, broadening to a blunt apex, papery, hirsute outside, glabrous inside, spreading, persistent, smaller than cluster. Staminate flowers 9 per cluster, shortly pedicellate; sepals granular glandular, variously hirsute; stamens 4, anthers 4-locular. Infructescences as staminate. Fruits 2 or 3 developing per cluster, young with ovary tomentose; pedicel pubescent, slender, reaching 12 mm; calyx hairy; styles subapical, 12, papillose, 3 mm, curved or straight, hairy calyx; maturing globose, 5 mm diam., stiffly leathery with dense short stout spines, tomentum and granular glands; calyx caducous; styles 5 mm, strap-like. Seeds globose 3mm diam., closely finely verrucose, with sarcotesta.
Distribution Papua New Guinea (Western Highlands, Sattelberg, Southern Highlands)
Habitat & Ecology Primary or secondary montane forests, 18002200m.
Note The mature fruits are like tiny durians. The doubly bullate leaves are noteworthy. Habit similar to M. carrii but kept separate by its more slender inflorescences.
Macaranga coggygria Airy Shaw, Kew Bull. 29 (1974) 324; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 143; Gen. Macaranga Prodr. (2008) 107. Type: Kairo NGF 45314 (holo K; iso L,SING), Papua New Guinea, Morobe Prov., Mumeng Subd., track to Mt. Shungol.
Small trees 6m. Twigs stout, 57 mm diam., terete, densely pubescent. Stipules oblong, 7 mm, apex acute, tomentellose without. Leaves: petioles 15 cm, stout, 3 mm diam., terete, densely pubescent, kneed at top; blades broadly ovate, 2330 by 1619cm, leathery, base broadly rounded to truncate, 13 cm peltate, margin with protruding glandular teeth, apex acute, sometimes with a 5 cm narrow caudex, drying khaki, glabrous above or with weakly pubescent main nerves, softly densely pubescent below and with sparse to fairly dense black granular glands, 3 main and several minor basal nerves. Staminate inflorescences with thread-like pubescent sinuous branches and 3 mm distant tiny flower clusters; bracteoles shortly stalked, spathulate, to 3 mm long, blade 1.5 mm, circular, outside hairy, inside entirely occupied by a patellar gland, spreading, persistent. Staminate flowers not seen. Infructescences a dense tangled mass, 14 cm across, ochreous pubescent panicles with 3 orders of strongly crooked intertwined axes, with spreading pale hairs; peduncle 3 cm; main axis stout, 2 mm diam., terete; bracteoles like staminate, mostly caducous. Fruits in groups of 13, globose, 3 mm diam., leathery, black granular glandular, with a few short caducous spines; pedicel thread-like, 814 mm, hirsute; calyx persistent, hirsute; style short, slightly lateral, 1 mm, papillose. Seeds flattened, coarsely tuberculate, with sarcotesta.
Distribution East New Guinea
Habitat & Ecology Disturbed forest and margins, montane forests, 12001900m.
Notes 1. Male inflorescence only known from a fragment.
2. The bracteoles are highly distinctive.
Macaranga congestiflora Merr., Philipp. J. Sc. Bot. 4 (1909) 282; Enum. Philipp. Flow. Pl. (1923) 440; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 310; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 34; Whitmore, Gen. Macaranga Prodr. (2008) 107. Syntypes: Curran 3590 (L); 3591 (K), both: Philippines, Palawan, near Puerto Princesa.
Small treelets to 6m. Twigs solid, angled, at first rusty scurfy. Stipules tiny, ovate, 2 mm, apex acute, papery, densely furfuraceous, erect, soon caducous. Leaves: petioles roundish, striate, to 14 cm, sparsely furfuraceous near tip; blades ovate, c. 15(24) by 13(22) cm, mostly very shallowly 3-lobed, papery to thinly leathery, base 3 cm peltate, upper margin with a few distant glandular teeth, apex very broadly rounded, drying either brown or slate grey, glabrous above, below with evanescent short white hairs on nerves and (very) sparse granular glands, sometimes weakly glaucous. Inflorescences small, congested, furfuraceous, axillary racemes; staminate crowded, to 45 in leaf axils, ultimately to 4 cm; main axis slender, rounded, scurfy, with a few distant, brittle 1015 mm branches and a few 5 mm tertiary branches, sometimes with axillary branches; bracts subpersistent, ovate 1.5 mm, hispid; flower clusters 3 mm, 35 crowded apically; enclosed by persistent, furfurescent, eglandular, papery, nerved bracteoles; latter ovate to oblong, 1.53 by 27 mm, conspicuous, apex divided to halfway or more into tapering, regular fimbriate strips. Staminate flowers 410 per cluster, tiny, 0.1 mm; pedicel very short, slender; sepals joined at base; stamens 3, anthers 4-locular. Fruits depressed globose, 8 mm across by 5 mm, weakly 6-shouldered, deeply trilobed, each lobe sulcate, leathery, drying blackish brown, smooth with fine granular glands except fissures, and evanescent sparse white hairs; pedicel to 720 mm; sepals small, persistent, free; stigmas persistent, tiny. Seeds 3, spherical, pitted.
Distribution Philippines (Palawan).
Habitat & Ecology Lowlands.
Macaranga conifera (Rchb.f. & Zoll.) Mόll.Arg. in DC., Prodr. 15, 2 (1866)vc1005; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 392; Whitmore, Tree Fl. Malaya 2 (1973) 112; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 157; Gard. Bull. Singapore 31 (1978) 53; in Airy Shaw, Kew Bull. 36 (1981) 321; Corner, Ways. Trees Malaya, ed.3 (1988) 300;. Whitmore, Gen. Macaranga Prodr. (2008) 109, Fig. 6. Mappa conifera Rchb.f. & Zoll., Linnaea 29 (1857) 466. Type: Teysmann s.n., 1857 (not found) Sumatra, prope Dermo Enim, Palembang.
Pachystemon populifolius Miq., Fl. Ind. Bat. Suppl. (1861) 462. Mappa populifolia (Miq.) Mόll.Arg., Linnaea 34 (1865) 198. Macaranga populifolia (Miq.) Mόll.Arg. (1866) 1006; Kurz, Forest Flora British Burma 2 (1877) 381; Hook.f., Fl. Brit. India 5 (1887) 450; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 322; Ridley, Fl. Malay. Penins. 3 (1924) 304; Corner, Ways. Trees Malaya (1940) 267.
Trees, sometimes reaching 30 m, dbh up to 60 cm; bole straight. Bark hooped, dippled, latex red; inner bark red; wood white. Twigs slender, to 4 mm diam., solid, terete, smooth, usually dark brown, finely furfuraceous becoming glabrous. Stipules lanceolate 5 by 2 mm, apex acuminate, dark brown, papery, finely furfuraceous, spreading, soon caducous. Leaves: petioles 3 cm, very slender, 1 mm diam., terete, glabrous; blades elliptic, occasionally trullate, c. 8(9) by 4(7) cm, membranous, base broadly rounded, to (almost) truncate (peltate in youth), margin entire, minutely recurved, apex acuminate, typically drying khaki, glabrous, closely below granular glandular and glaucous, secondary nerves very delicate often darker than lamina, sometimes slightly raised below. Staminate inflorescences amongst and behind leaves, crowded, dense, glabrous, 34 cm racemes (or with a few short third order axes); main axis terete; peduncle 520 mm, granular glandular, slender; branches alternate, catkin-like, with dense overlapping bracteoles enclosing flower clusters; bracts entire, oblong, 2 mm, soon caducous; bracteoles deltoid, 3 by 2 mm, apex acute, coarsely regular dentate, papery, glabrous, both sides granular glandular, persistent, erect. Staminate flowers in clusters of c. 12, tiny, 0.2 mm; pedicel 0.2 mm, glabrous; sepals free; stamen 1, anther 4-locular. Infructescences as staminate. Fruits crowded, bilobed, 4 by 8 by 4 mm, woody, closely yellow brown granular glandular; pedicel slender 2 mm; calyx minute, papery, unlobed, reflexed; stigmas 2, persistent, apical, short, recurved, smooth. Seeds with a thin sarcotesta, globose, testa with flat warts.
Distribution India (Andaman Islands; Kurz 1877), Peninsular Malaysia, Sumatra, Borneo, Sulawesi (Malili).
Habitat & Ecology Slik: In disturbed places in primary forest (gaps) and in secondary forest, including scrub vegetation. Along streams, roads, skid trails. Dry to periodically inundated to marshy terrain. Loam, sandy, and limestone soils. Up to 1100 m altitude.
Notes 1. The very thin finely dark-nerved leaves readily distinguish this from its relative M. amissa.
2. Kostermans 12037 from Sumatra, Palembang, differs in possession of glands on the leaf base near the petiole insertion.
3. The Sulawesi occurrence (Ota s.n., 21/4/32, L!, and Ota s.n., 25/4/32, L!) is somewhat surprising. Apart from these collections the whole Conifera species group is confined to the Sunda Shelf.
4. Slik: This species has a very strong dimorphy between the juveniles and the adults. The juveniles have bright red petioles, clearly peltate leaves and conspicuously large green persistent stipules.
Macaranga constricta Whitmore & Airy Shaw, Kew Bull. 25 (1971) 238; S.J. Davies, Harvard Pap. Bot. 6 (2001) 395. Type: Whitmore 3156 (holotype: K; isotypes: A, KEP, L, SING), Malay Peninsula, Pahang, northeast Gunong Benom, Ulu Krau drainage.
Tree, glaucous, to 27 m tall, c. 30 cm dbh; twigs 610 mm in diam., distinctly constricted at the nodes and swollen at the internodes, densely glaucous, glabrous or minutely ferrugineus pubescent at the apex, hollow, housing ants. Bark smooth, hoop-marked, grey. Stipules ovate, 58 by 57 mm, the pair not completely encircling the twig, recurved, succulent, abaxial surface producing food-bodies, 13 pairs present on the shoots. Leaves: petioles terete, 1020 cm long, densely glaucous, glabrous; blades ovate, rather consistent in size and shape, 1528 by 1020 cm, shallowly trilobed, dissected to 1/4 to 1/3 of the leaf length, central leaf lobe 58 cm wide at the base, lateral lobes ascending 36 cm wide, 26 cm peltate, base narrowly rounded, margin entire, without small protruding nectaries, apices acuminate, narrowing quite suddenly, adaxial surface glabrous, abaxial surface with minute scattered hairs on the veins and otherwise mostly glabrous, very densely glaucous; 1° venation palmate with 67 prominent veins, 2°4° venation feint, 2° venation scalariform, strongly looping near the leaf margins and joining with the adjacent veins and not meeting the leaf margin; young leaves glaucous, with prominent yellow granular glands on abaxial surface, minutely ferrugineus pubescent on the veins. Staminate inflorescences paniculate, erect, long and spreading, 1828 by 1218 cm, axes mostly glabrous and densely glaucous, distal branches scattered with minute ferrugineus hairs, to 4 axis orders, main axis flattened c. 1/31/2 of total inflorescence length, first pair of secondary branches ± opposite with accessory branches; bracts 57 by 23 mm, persistent, margin entire, apex acute, glabrous or scattered with ferrugineus hairs; flower clusters with c. 2025 flowers, spirally arranged and clustered on ultimate inflorescence branches; bracteoles ovate-elliptic, 57 by 34 mm, erect, enclosing flower clusters, margin entire or rarely slightly lobed, apex narrowly acuminate to caudate and terminating in a fine conical gland, glabrous or rarely scattered with minute red-brown hairs towards the base, scattered with yellow granular glands, adaxial surface with a dense patch of red-brown hairs at the base. Staminate flowers 11.5 mm long, pedicellate; sepals free, apex scattered with minute red-brown hairs; stamens (2)3(4); anthers 3- (or 4-)locular. Pistillate inflorescences shortly paniculate, erect, 514 by 410 cm, stout, minutely ferrugineus pubescent, basal axes glaucous and less densely pubescent, up to 3 axis orders, secondary branches ±opposite without accessory branches; bracts elliptic, 45.5 by c. 2 mm, apex acute, often terminated by a minute conical gland, glaucous, scattered with minute ferrugineus hairs particularly on the margins and towards the base, caducous. Pistillate flowers 35 mm long, solitary in bract axils; calyx urceolate, c. 2 mm long, apex truncate to very slightly 4-lobed, scattered with fine red-brown hairs, glaucous, persistent; ovary 4-carpellate, c. 2 mm long; styles c. 3 mm long, fused for c. 4/5 of the length into a prominent column, free and recurved only at the apex, persisting to form a prominent 34 mm long crown at fruit apex; stigma not dissected. Fruits subglobose, 911 by 57 mm, pedicellate, with one discrete glandular patch on each carpel wall developing into a slightly raised disc covered in yellow, sticky exudate, glaucous; pedicel 46 mm long, glabrous or with scattered minute ferrugineus hairs, glaucous. Seeds subtriangular-ovoid, 44.5 mm in diam., black, with shallow coarse grooves and a small cruciform scar, encased in a fleshy bright red aril.
Distribution Malay Peninsula (endemic in Pahang and Trengganu).
Habitat & Ecology An early successional tree. Numerous collections have been made on Gunong Benom. Macaranga constricta grows to be quite a large early successional tree. It occurs on hillsides and ridges in lowland forest to 900 m.
Note The densely glaucous leaves and twigs, and gross flower and fruit morphology are reminiscent of the closely related M. griffithiana. It differs from this species in having prominently constricted twig nodes, leaf 2° veins looping very strongly and not connecting with the leaf margins, an absence of conical nectaries on the leaf margin, longer staminate inflorescences with fewer well-spaced branches that protrude above the shallow canopy, staminate bracteoles with entire margins and a much longer acumen, leaf lobes more narrowly ascending, and the pistillate flowers with much longer connate styles. In addition, M. constricta occurs in dry-land forests whereas M. griffithiana occurs primarily in fresh water swamps and flooded areas.
Macaranga costulata Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 338; Merr., Bibl. Enum. Born. Pl. (1921) 341; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 156; Gen. Macaranga Prodr. (2008) 112, Fig. 7. Type: Haviland & Hose 3662 (iso K, L, P, SING), Borneo, Sarawak.
Trees, usually only 5 m (Slik: trees up to 30 m tall, dbh up to 60 cm; latex red). Twigs solid, slender, to 4 mm diam., terete, weakly furfurescent. Stipules broadly elliptic, to 10 by 2mm, papery, pubescent, erect, soon caducous. Leaves: petioles c. 6 cm, slender, terete, finely ridged, (almost) glabrous, not kneed; blades ovate-oblong to trullate, 10(15) by 47(12) cm, papery, base broadly truncate occasionally 13 mm peltate, rarely cordulate, 2 large conspicuous glands at petiole insertion, margin entire sometimes very weakly crenate rarely shallowly dentate, apex acute to abruptly acuminate, usually drying pale greenish khaki, below sparsely pubescent on main nerves. Staminate inflorescences crowded, broad fairly dense delicate panicles to 8 by 5 cm, 3 axis orders, finely furfuraceous to almost glabrous; main branches divaricate; peduncle short, 310 mm; main axis slender. under 1 mm diam., roundish, coarsely striate; lower branches alternate, axillary branches very rare; bracts not seen; branches thread-like, tertiary branches arising from flower clusters on secondaries; flower clusters evenly spaced, distant, 510 mm apart; bracteoles linear, 34 mm, distorted by 13 patellar glands, or pointed, occasionally with a few marginal teeth, glabrous, base spathaceous, enclosing cluster, spreading, conspicuous, persistent. Staminate flowers c. 15 per cluster, developing sequentially, 0.5 mm diam., sessile; sepals 3, glabrous; stamens 3, anthers 4-locular. Pistillate inflorescences crowded, delicate, diffuse narrow racemes or spikes, to 10 cm long, as staminate. Fruits in groups of 13, depressed globose, slightly flattened, weakly bilobed, 2 by 3 mm, finely golden granular glandular; pedicel thread-like, 1 cm; calyx tiny, persistent; styles 2, short, recurved, apical. Seeds with sarcotesta, shallowly coarsely rugose.
Distribution Borneo (Sarawak, Sabah, Kalimantan east and central).
Habitat & Ecology Whitmore & Slik: In primary and secondary forest, including scrub. On clay, dark brown, limestone and sandstone soils. Lowlands to 1800m.
Note The fruits of Kostermans 6026, a big tree from the Kutai limestone, have a few tiny spines.
Macaranga crassistipulosa Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv, add. VI (1919) 26; Airy Shaw, Kew Bull. 37 (1982) 26; Whitmore, Gen. Macaranga Prodr. (2008) 114. Type: Warburg 15570 (not seen), Sulawesi, Minahassa.
Juvenile parts subglabrous. Twigs slender. Stipules 56 mm long, apex acute, persistent, thick, glabrous. Leaves: petioles 35 cm; blades oblong or ovate-oblong, 712 by 2.56 cm, base narrowly cordate, above with 2 big glands, margin unevenly finely toothed, apex cuspido-acuminate, shiny above, below dull and slightly pruinose, below densely granular-glandular, glabrous, penninerved, base shortly 3-nerved, secondary nerves c. 10. Staminate inflorescences a panicle 67 mm [?cm] long, divaricate-racemose, stipe short, glabrous; rachis compressed; bracteoles triangular-semiorbicular, 12 mm long, apex obtuse or acute, glabrous, 59-flowered. Staminate flowers with calyx scarcely granular-glandular, opening 1 mm wide, stamens 3 or 4, anthers 4-locular. Pistillate flowers, fruits and seeds unknown.
Distribution North Sulawesi (known from the type only).
Note Not seen by me. Translated from Pax & Hoffmann (1919). The description suggests that this is a member of the Angustifolia group.
Macaranga cucullata J.J.Sm., Nova Guinea Bot. 8 (1910) 237, t.60; Nova Guinea Bot. (1912) 789; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 369; L.M.Perry, J. Arnold Arb. 34 (1953) 202; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 144; Gen. Macaranga Prodr. (2008) 114. Type: Versteeg 1066 (not seen; perhaps mistake for Versteeg 1064: K, L, P, SING), Indonesia, Papua, Noordfluss.
Trees. Twigs ? Stipules, Leaves like M. aleuritoides except leaves with quaternary nerves reticulate, somewhat papery, and with the minor nerves below not strongly raised and puberulous. Stipules united, single. Staminate inflorescences panicles like in M. aleuritoides but glabrous and bracteoles strongly cucullate, 1.5 by 1.5 mm, with a broad rounded 1 mm protruding tip, bearing 1 large gland, glabrous, margins entire, spreading, larger than clusters; latter closely evenly spaced along all axis tips. Staminate flowers 6 per cluster with long pedicel, stamens 3 or 4, anthers 2-locular. Infructescences 33.5 cm. Fruits few, depressed squarish globose, densely glandular with pubescent 2 mm spines.
Distribution New Guinea
Habitat & Ecology Primary and secondary lowland forest.
Note Very similar to the much commoner M. aleuritoides but quite distinct in both leaf details and the male bracteoles and flowers. Fruit description taken from Perry (1953).
Macaranga cumingii Mόll.Arg. (1866) 1005; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 342; Merr., Enum. Philipp. Flow. Pl. (1923) 441; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 34; Whitmore, Gen. Macaranga Prodr. (2008) 115. [Adenoceras cumingii Baill., Ιtude Euphorb. (1858) 430, nomen nudum.] Type: Cuming 1100 (iso: CGE, K, OXF, P, W), Philippines, Luzon, Albay Prov.
Macaranga caudatifolia Elmer, Leafl. Philipp. Bot. 2 (1908) 427; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 342; Merr., Enum. Philipp. Flow. Pl. (1923) 440; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 34.
Macaranga cuneata Elmer, Leafl. Philipp. Bot. 2 (1908) 428.
Macaranga sibuyanensis Elmer, Leafl. Philipp. Bot. 3 (1910) 922.
Macaranga apoensis Elmer, Leafl. Philipp. Bot. 7 (1915) 2646; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv, add.VI (1919) 27.
Small trees to 10m. Twigs slender, rounded, finely to sparsely furfuraceous becoming glabrous. Stipules elliptic, 4 mm, furfuraceous, erect, very soon caducous. Leaves: petioles 27 cm, slender terete, finely furfuraceous to glabrous; blades ovate to elliptic, 812 by 36 cm, papery, base cuneate to truncate, sometimes with 12 glands at petiole insertion, apex acuminate to caudate, usually glabrous, occasionally hairy on nerves below, lower surface with fine patchy granular glands, secondary nerves c.7 pairs. Staminate inflorescences crowded, 58 cm spikes or occasionally with a few short branches; axes slender, furfuraceous; flower clusters small, evenly spaced; bracteoles inconspicuous, size and shape variable, ovate, 13(7) mm long, base sometimes narrowed, apex acute or (long) acuminate, with 24 big adaxial glands, glabrous, soon caducous or a few persisting. Staminate flowers c. 16 per cluster, 0.08 mm diam.; pedicel long; calyx divided to base; stamens 3, anther 2-locular. Pistillate inflorescences: crowded, spike, 613 cm, axes slender, furfuraceous becoming glabrous; flower clusters 12 cm distant, mainly or wholly on distal half; bracteole as staminate. Pistillate flowers very long pedicellate, c. 3 per cluster; calyx entire or divided to halfway; ovary with rugose globose eccentric stigma. Fruits clustered 23(5), globose, 45 mm, smooth, thinly woody, at first with a waxy coating, sometimes with warty excrescence; pedicel thread-like, very long, 1530 mm; calyx small, persistent; stigma a small rounded sessile slightly eccentric plate, caducous. Seeds 1, round, verrucose.
Distribution Philippines (Luzon, Biliran, Sibuyan, Negros, Panay, Mindoro, Samar, Mindanao).
Habitat & Ecology 10002200m.
Notes 1. Mόller (1866) took up Baillon's (1858) nomen nudum, used the same number (Cuming 1100) as type, and treated it as a basionym. The mistake has been perpetuated ever since. Elmer subsequently described three species in 19081910, all considered to be M. caudatifolia by Pax & Hoffman (1914) and another in 1915 reduced by Merrill in 1923.
2. I believe only one species is involved in this whole complex, for which M. cumingii is the earliest name. The bracteole, which is usually such a reliable diagnostic character, is very variable in size and shape (even on the single specimen Elmer 10719, type of M. apoensis). Leaf apex is variable and occasionally strongly caudate. The plant is usually glabrous, but occasionally at first pubescent to furfuraceous. Fruit ornamentation is variable. The most distinctive characters are the very long female pedicel, especially in fruit, clustered fruits, with small plate-like slightly eccentric stigmas, and flower clusters borne mainly in the distal part of the narrow slender inflorescences. The species has a wide range in the mountains. Study of the living tree will either enable discrete segregate species to be recognised all as parts of a tight complex, or confirm my conclusions that the characters that have been used to distinguish species are in fact not constant within a tree or population. See M. sylvatica, another variable species from the Philippines.
Macaranga darbyshirei Airy Shaw, Kew Bull. 29 (1974) 323; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 144; Gen. Macaranga Prodr. (2008) 118. Type: Darbyshire 396 (holo K; iso L), Papua New Guinea, Sepik Dist., Lumi Subdist., near Wigote village (Torricelli Mountains).
Trees to 20m. Twigs fairly slender to stout, 48(15) mm diam., angled, usually strongly glaucous, glabrous. Stipules elongate-triangular, c. 25 by 8 mm, with straight sides tapering regularly from base to a long narrow acuminate apex, both margins revolute so apparently only 2/3 this width, chestnut-brown but strongly whitish glaucous, glabrous except for midline, stiffly papery, usually recurved, usually rather straight, persistent. Leaves: petioles 917(20) cm, slightly flattened, 24 mm diam., striate, (strongly) glaucous, glabrous or weakly pubescent; blades ovate to ovate oblong, 1520(40) by 814(37) cm, leathery, base rounded (sometimes deeply so, with blade widest at middle), deeply 35(11) cm peltate, occasionally with flat glands on main nerves midway between margin and petiole insertion, margin repand, sometimes with inconspicuous distant small protruding glands, apex 2 cm, abruptly caudate, drying khaki below often with the main nerves darker, below often greyish glaucous, above maturing pubescent just on main nerves, below weakly velvety from pubescent nerves and with close small golden granular glands, spider's web nervation with tertiaries and faint quaternaries scalariform. Staminate inflorescences open panicles to 26 by 10 cm, axes distally furfuraceous, glaucous; main axis flattened, 34 mm diam., sometimes striate; peduncle c. 8 cm; secondary axes to 10 cm, normal to main axis, lower ones 4 cm spaced, tertiaries ascending, to 3 cm, with some axillary branches, a few branches arising from a flower cluster; bracts soon caducous, ovate, 12 by 5 mm, apex acute, warm brown, papery; bracteoles ovate, to 4 by 3 mm, apex shortly cuspidate, strongly cucullate, nerved, margin entire or coarsely irregularly crenate or pectinate dentate, densely finely pubescent, loosely enclosing irregularly spaced to contiguous flower clusters. Staminate flowers in clusters of c. 20, shortly pedicellate with 46 stamens with anthers 4-locular. Infructescences short, 611 cm, stout golden pubescent spikes, axis flattish, 3 mm diam.; peduncle 4 cm. Fruits clustered at apex, globular or weakly bilobed, c. 8 by 10 mm, woody, with sinuous 5 mm sometimes shortly branched spines, dense or diffuse, these and wall (closely) golden granular glandular; pedicel stout, 510 mm; calyx caducous. Seeds globose, 6 mm, coarsely irregularly rugose, with sarcotesta.
Distribution New Guinea.
Habitat & Ecology Lowlands to 900m, primary or secondary forest.
Macaranga daviesii Takeuchi, Harvard Pap. Bot. cxc (2007) 389, fig. 14. Type: Takeuchi, Towati & Ama 19931 (holo LAE; iso A, BISH, BO, BRI, CANB, K, L, MO), Papua New Guinea, Enga Prov., Paiela Bivouac (Lake Tawa).
Subcanopy trees, 1520 m tall; bole straight, erect, unbuttressed; crown numerously branched. Wood straw, dense. Twigs terete, subapical diam. (2)2.53.5(4) mm, firm, not pithy, resinous, exudate black, vernicose, obscurely striate, elenticellate; internodes (0.5)14(5.5) cm. Indumentum acicular hairs, 23 mm long, appressed (or patent), easily dislodged with handling, irritant, dense, persisting, orange-brown (fresh pink on all vegetative parts); scale glands 0.10.2 mm diam., hyaline-yellow or pellucid. Stipules united, apparently single, sheathing, (32)4062(76) by 610 mm, terminal, erect, caducous, indument as twigs. Leaves: petioles (7)1535(42) by 13 mm, planoconvex, pulvinate at both ends or not, black, appressed hairy; blades ovate-rhomboid, (4.5)713(15.5) by (2.5)610.5(12.5) cm, rugulose, coriaceous, base broadly rounded, notched 24 mm from the petiole insertion, obscurely auriculiform-lobulate, auricles marked by 1 (or 2) petelliform glands, margin crenate or eroded, often reflexed, apex obtuse, upper surface very dark green, dried dull grey (or burnneous), glabrescent, lower surface pruinose, dried orange-brown, strigose on major vines, lanulose or villous on tertiary veins, domatia swollen, foveate or not, black, obscured by hair tufts; venation usually craspedrome, secondaries in 58 pairs above a basal digitate nervation, straight-divergins 2540Ί from costae, tertiary nerves scalariform, all veins deeply impressed above, prominent below, reticulum tessellate, regular, areoles abaxially white-papillate, densely furnished with yellow granules. Staminate unknown. Pistillate flowers solitary, axillary, erect; peduncles cylindrical, (20)2436(40) by 0.81.2 mm, black, strigose, granulose-glandular; bracts 4 in 2 dimorphic series, contiguous with calyces; major (outer) bracts: stipitate, trullate, (7)1013 by 35 mm inclusive a 24 mm stipe, fugacious, crenate, ascending, costate, induplicative, fleshy, adaxially patelliform-pluriglandular, abaxially with indument as the peduncle; minor (inner) bracts: inserted 90Ί from the larger pair, ovate-deltoid, (2)3 by 23 mm, persisting in fruit, hirtellous; calyx globose-ovoid, 45 mm diam., completely enclosing the ovary except for the style-ostiole, chartaceous, rupturing irregularly at anthesis, decidous, outer surfaces with indument as the peduncle, inner surfaces smooth; ovary ovoid, c. 4 by 3.54 mm, scurfy, laxly hirtellous, bilocular; styles 2, (5)79 mm long, arcuate, plumosely lobed. Fruits inserted on accrescent peduncles to 65 by 2 mm, globose or ovoid, 913 by 1014 mm (immature); exocarp jet black, encircled by a longitudinal suture, glandular-scurfy, excrescences spiniform, subulate, 1 mm long, laxly pilosulous. Seeds ovoid, c. 6 by 5 mm, muricated, crustateceous, brunnescent.
Distribution Papua New Guinea (known from type locality only).
Habitat & Ecology Mossy montane forest on limestone karst, a mature-phase entrant in forest community development. Flowering and fruiting: August. Altitude: 2600 m.
Notes 1. With long-peduncled solitary flowers and conspicuously connate stipules, M. daviesii is obviously assignable to the Longistipulata group.
2. In M. daviesii the anthetic female calyx is metreform, completely enclosing the ovary except for a small orifice at the summit through wich the style arms are exserted. This is similar in M. eymae. During flower maturation the calyx tears basipetally and appears apert or lobed.
Macaranga decipiens L.M.Perry, J. Arnold Arb. 34 (1953) 212; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 144; Gen. Macaranga Prodr. (2008) 118. Type: Kanehira & Hatusima 13256 (holo A; iso BO), Indonesia, Papua, Vogelkop, Manokwari, Waren.
Shrubs or treelets, or once a 22 m tree. Twigs slender, 3 mm diam., angled, tomentose, sometimes with long curved internodes suggesting sarmentose habit. Stipules elliptic, 10 by 1 mm, hirsute, erect, persistent. Leaves: petioles 23 cm, slender, 1.5 mm diam., terete, pubescent, kneed at both ends; blades ovate-elliptic, c. 16 by 5 cm, papery, narrowing to rounded base, shallowly cordulate, sometimes with 2 inconspicuous marginal glands near petiole insertion, apex narrowing to a 15 mm caudex, drying darkish, tomentose on lower midrib above, and sparsely with spreading white hairs on nerves below, only sparsely patchily granular glandular below. Staminate inflorescences more or less adpressed to stem, short, 5 cm, hirsute racemes, with a few erect branches; peduncle 1 cm; main axis flattened, 2 mm diam., with spreading white hairs;, branches alternate, some upper with short axillary branches; flower clusters close evenly spaced; bracteoles spreading longer than clusters, slightly fleshy, 2 by 0.5 mm with trapezoid, acute head tapering downwards, 23 small patellar glands inside, granular glandular both sides, margin entire, sparsely hairy. Staminate flowers in clusters of c. 6, 0.3 mm diam., shortly, stalked; sepals 3; stamens 23, anthers 4-locular. Pistillate flowers calyx not seen; ovary 2-locular, style 35 mm, plumose. Infructescences spikes, 313 cm, axis slender, slightly flattened, tomentose with spreading white hairs, with a usually single terminal fruit subtended by several bracteoles, latter elliptic, 5 by 1 mm, hirsute, acute, with adaxial patellar glands. Fruits usually solitary, globose or bilobed, 6 by 9 mm, closely invested with short 12 mm sinuous hairy spines, ripening red, sessile. Seeds unknown.
Distribution Indonesian New Guinea.
Habitat & Ecology Lowlands. Primary and secondary forests.
Note Pistillate flower description taken from Perry (1953). The red fruit is diagnostic.
Macaranga densiflora Warb., Bot. Jahrb. 13(1891) 350; K.Schum., Notizbl. Kφn. Bot. Gart. Berlin 2 (1898) 128; K.Schum.& Lauterb., Fl. Schutzgeb. Sόdsee (1900) 396; J.J.Sm., Nova Guinea Bot. 8 (1910) 238; Nova Guinea Bot. 8 (1912) 791, Nova Guinea Bot. 12 (1917) 546; Rechinger, Denkschr. Akad. Wiss. Wien 89 (1913) 568; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 372, t.62AE; in Engl., Pflanzenr. IV.147.xvii, add. VII (1924) 185; Kaneh. & Hatus., Bot. Mag. Tokyo 52 (1938) 411; L.M.Perry, J. Arnold Arb. 34 (1953) 219; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 144; Gen. Macaranga Prodr. (2008) 119. Type: Warburg probably 20509 (not seen), Papua New Guinea, Madang Pov., Hatzfeldthafen.
[Macaranga tiliacea Rόbsaamen, Marcellia 4 (1905) 15, nomen nudum, see Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xvii, add. VII (1924) 185.]
Macaranga acuminata Ridl., Trans. Linn. Soc. Bot. 9 (1916) 148.
Small trees to 5 m. Twigs slender, 3 mm diam., terete, finely pubescent to almost glabrous. Stipules elliptic, 5 by <1 mm, apex acuminate, papery, pubescent, erect, soon caducous. Leaves: petioles 1.516 cm, slender, 1.5 mm diam., terete, finely pubescent, kneed at top; blades very variable, elliptic, 10 by 3.5 cm, or less usually trapezoid, 12(21) by 6(11) cm, papery, base (subtruncate to) broadly cuneate, often with 2 glands sunken above and raised below at petiole insertion, occasionally slightly protruding beyond margin, margin repand and with tiny distant protruding glandular teeth, apex narrowing to a 5 cm caudex, usually drying grey brown above and darker below, below hairy on midrib and to a lesser extent all other nerves, closely granular glandular on lamina and sometimes somewhat on main nerves, above finely pubescent on main nerves and lamina scabrid, 3 nerves at petiole insertion, the laterals running close to margin half way up lamina, secondary nerves rather (1 cm) distant. Staminate inflorescences delicate, open, finely pubescent racemes', 49 cm long; branches ascending to divaricate, thread-like, to 3 cm long; peduncle 2.54 cm; main axis slender, 1 mm diam., terete; lowermost branches (sub)opposite; bracts tiny, elliptic, 1 mm long, apex acuminate, pubescent outside, erect, persistent; flower clusters contiguous in loose catkin-like clusters, 5 by 38 by 5 mm at secondary branch tips and crowded tightly on 5 mm tertiary branches, subtended by bracteoles; latter trilobed 1 by 1 mm, spreading, thick, the lobes narrow, spreading, granular glandular outside, inside glabrous and bearing 2 small patellar glands. Staminate flowers 45 per cluster, subsessile; stamens 2, anthers 4-locular. Infructescences spikes, c. 9 cm lon, axis slender, 1 mm diam., terete, finely pubescent, with a terminal and sometimes a few subterminal clusters of fruits subtended by leafy bracts; latter ovate 5 by 315 by 10 mm (smaller in flower) , acute, somewhat cucullate, margins deeply finely toothed, both sides pubescent, pinnately nerved, sometimes with stalk to 3 mm. Fruits 12 per cluster, round, 5 mm diam., sessile, thinly woody, yellow velvety and with sparse black granular glands and caducous soft stout 1 mm velvety spines; calyx persistent; style lateral, strap-like, 3 by 1 mm, pectinate on outer side, persistent. Seeds round, coarsely verrucose, with sarcotesta.
Distribution Throughout New Guinea, the Bismarck Archipelago, Solomon Islands.
Habitat & Ecology Riversides and secondary forest. Lowlands, once at 450 m and 950 m.
Note A variable species. The staminate flower clusters in loose terminal compound catkins are distinctive, as are the leaf base glands, which however are sometimes absent. That the catkins contain branches, albeit short, stictly makes the staminate inflorescence a 3 axis-order panicle, not a raceme.
Macaranga denticulata (Blume) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1000; Kurz, For. Fl. Brit. Burma 2 (1877) 387; Hook.f., Fl. Brit. India 5 (1887) 446; Brandis, Indian Trees (1906) 591; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 334, t. 54B; Ridl., Fl. Malay. Penins. 3 (1924) 303; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 446; Corner, Ways. Trees Malaya (1940) 265; Backer & Bakh.f., Fl. Java 1 (1963) 488; Airy Shaw, Kew Bull. 26 (1972) 288; Whitmore, Tree Fl. Malaya 2 (1973) 111; Gard. Bull. Singapore 31 (1978) 53; Chiu, Guihaia 2 (1982)148, in key; Long, Fl. Bhutan 1 (1987) 804; Chakrab. & Gangop., J. Econ. Taxon. Bot. 13 (1989) 597; Chakrab. & N.P.Balakr., J. Econ. Taxon. Bot., Addit. Ser. 9(1992) 80; Gardner, Sidisunthorn & Anusarnunthorn, For. Trees N. Thailand (2000) 312; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 364, Plate XX: 1; Whitmore, Gen. Macaranga Prodr. (2008) 120. Mappa denticulata Blume, Bijdr. (1825) 625; Reichb.f. & Zoll., Verhand. Natuurk. Vereen. Ned. Ind. 1(1856) 9; Linnaea 28 (1856) 308. Type: Blume s.n. (holo L?; iso K, P, W), Java, in silvis montium Salak et Gede.
Rottlera glauca Hassk., Flora 25, Beibl. 2 (1842) 41.
Mappa gummiflua Miq., Fl. Ind. Bat. Suppl. (1858) 458. Macaranga gummiflua (Miq.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1000.
[Mappa wallichii Baill., Ιtude Euphorb. (1858) 430, nomen nudum.]
Mappa chatiniana Baill., Adansonia 1 (1861) 349 (chantiniana). Macaranga chatiniana (Baill.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 996; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 318.
Mappa truncata Mόll.Arg., Linnaea 34 (1865) 198.
Macaranga denticulata (Blume) Mόll.Arg. var. zollingeri Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1000.
Macaranga perakensis Hook.f., Fl. Brit. India 5 (1887) 447.
Macaranga henricorum Hemsley, J. Linn. Soc. 26 (1894) 442; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 446.
Macaranga denticulata (Blume) Mόll.Arg. var. pustulata (King ex Hook.f.) Chakrab. & Gangop., J. Econ. Taxon. Bot. 13 (1989) 597.
Small trees. Twigs to 8 mm diam., strongly angled and ridged, densely fawn furfuraceous. Stipules elliptic, 36(10) by 12(3) mm, densely fawn furfuraceous, leathery, erect, very soon caducous. Leaves: petioles 511 cm, fairly slender, round, thinly fawn furfuraceous, kneed at top; blades deltoid (10)15(24) by (8)13(18) cm, thinly leathery, base 1 cm peltate (with several small inconspicuous glands at petiole insertion) broadly truncate or broadly slightly rounded, margin entire, apex acute, sometimes acuminate, drying often olive above, fawn below, nerves below finely furfuraceous to pubescent becoming glabrous, secondary nerves straight and parallel, 3 main nerves from the petiole insertion. Staminate inflorescences dense overlapping small panicles, to 9 by (6)9 cm, 3(4) axis orders, axes fawn furfuraceous; branches all divaricate; peduncle 1015 mm, 1 mm diam., ridged; lowest branches mostly (sub)opposite, others alternate, without axillary branches, ultimate branches 1 cm, thread-like; bracts tiny, inconspicuous, 1 mm, rounded, furfuraceous outside, persistent, erect; flower clusters small, 2 mm across, evenly spaced, at maturity not touching; bracteoles as bracts, shorter than clusters. Staminate flowers globose, developing in sequence, becoming 1 mm diam., sessile, c. 6 per cluster; calyx divided to base, granular glandular near apex; stamens c. 15, 4-locular. Pistillate inflorescences dense, overlapping small racemes, to 7 by 3(5) cm, 2 axis orders, fawn furfuraceous; branches divaricate, round to slightly flattened; peduncle smooth or slightly striate, to 15 mm, lowest branches subopposite or alternate; bracts and bracteoles as staminate. Fruits 1(3) per cluster, divaricate, strongly bilobed, broader than long, 3 by 5 mm, smooth blackish brown, wholly finely dark granular glandular; pedicel stout, to 1 cm; calyx small, persistent; styles 2, short, finely papillos, apical. Seeds round, with sarcotesta, shallowly verrucose.
Distribution India (Assam, Sikkim, Andaman Islands; Chakrabarty & Balakrishnan, 1992), Nepal, Bhutan, Sri Lanka, China (Yunnan, Hainan), Laos, Vietnam, Cambodia, Thailand, Malay Peninsula (mainly north), Sumatra, Java.
Habitat & Ecology Common in secondary forest. Sea level to 1200m.
Notes 1. Flowers and fruits similar to M. pustulata.
2. There is no doubt that M. chatiniana, based on a plant cultivated in the Paris Botanic Garden hothouses (Chatin s.n.,1856, P!, W!), is M. denticulata.
Macaranga depressa (Mόll.Arg.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 989; Whitmore, Kew Bull. 29 (1974) 445; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 146; S.J. Davies, Harvard Pap. Bot. 6 (2001) 396. Pachystemon depressus Mόll.Arg., Flora 47 (1864) 465. Type: See forms.
Small tree to 10 m tall, but often smaller or shrub-like; twigs slender, 37(8) mm in diam., solid, not housing ants. Bark smooth, pale grey-brown. Stipules deltate, 612 by 511 mm, the pair not completely encircling the twig, leathery, not succulent, spreading and usually perpendicular to the twig, usually 48 or more pairs present on the shoots. Leaves: petioles terete, slender, 820 cm long, not glaucous; blades vary among forms. Staminate inflorescences paniculate, erect, to 25 by 10 cm, drying black, to 3 axis orders, main axis flattened, first pair of secondary branches ± opposite with accessory branches; bracts ovate-elliptic, 514 by 47 mm, margin entire, apex acute, caducous; flower clusters with 2040 flowers, spirally arranged and grouped towards the ends of inflorescence branches, mostly much larger than the bracteoles; bracteoles small, broadly ovate, 34 by c. 3 mm, only partially enclosing flower clusters, margin entire, apex acute to shortly acuminate. Staminate flowers c. 1 mm long, sessile; sepals fused, splitting irregularly to c. 1/2, apex densely covered in minute red-brown hairs; stamens 1; anthers 3-locular. Pistillate inflorescences paniculate, erect, 412 by 46 cm, stout, either unbranched or with one pair of short opposite branches without accessory branches, the flowers clustered at ends of the axes; bracts ovate, c. 5 by c. 4 mm, apex acute to acuminate, caducous. Pistillate flowers 35 mm long, solitary in bract axils; calyx urceolate, 12 mm long, densely ferrugineus pubescent, with scattered yellow granular glands towards the apex, persistent, splitting irregularly as ovary expands; ovary 45-carpellate, c. 1.5 mm long; styles 23 mm long, with long and finely acuminate apices, expanding as fruit matures, fused at base, free and spreading from c. 1/3 of the length, persisting to form a prominent 35 mm long crown at fruit apex; stigma not dissected. Fruits subglobose, compressed, 68 by 811 mm, sessile, one discrete glandular patch on each carpel wall developing into a long slender horn-like process 37 mm long, covered in yellow, sticky exudate, glabrous or rarely with some scattered ferrugineus hairs towards the base and on the sutures. Seeds subtriangular-ovoid, c. 4 mm diameter, black, with shallow coarse grooves and a small cruciform scar at the columella attachment point, completely encased in a fleshy bright red aril.
Notes 1. Distinguishing characteristics, and ecological and distributional information for each of the forms of M. depressa are listed in their descriptions below.
2. Macaranga depressa is very closely related to M. triloba. Both species are in the core group of Macaranga section Pachystemon. They differ from other members of the section in having solid twigs that are not inhabited by ants, spreading coriaceous stipules, and dense sharp erect hairs on the young leaf bud. Macaranga depressa differs from M. triloba in having smaller staminate bracteoles that do not completely surround the flower cluster, staminate bracteoles with shortly acute and less elongate apices, more slender and delicate staminate inflorescence axes, less stout twigs, and usually smaller and less prominently dentate leaves.
3. Both M. depressa and M. triloba have very broad ecological distributions. All three forms of M. depressa and M. triloba are known from the lowlands to above 1200 m. Macaranga depressa forma strigosa is the most distinct of these taxa due to the strigose hairs on the twigs and leaves, and in submontane areas it often has subentire leaves. In the Kelabit Highlands of Sarawak Davies (2001) has seen M. depressa forma strigosa and forma glabra growing sympatrically. The other form of M. depressa does not overlap in distribution, and some specimens are particularly difficult to distinguish from M. triloba. All three of these taxa show wide morphological variation. An analysis of morphological and molecular variation across the full range of these taxa would be extremely useful in clarifying their status. Such an analysis may reveal a single widespread and polymorphic taxon.
a. f. depressa
Macaranga depressa (Mόll.Arg.) Mόll.Arg. f. depressa: S.J. Davies, Harvard Pap. Bot. 6 (2001) 397. Macaranga depressa (Mόll.Arg.) Mόll.Arg. var. genuina Mόll.Arg. in DC., Prodr. 15, 2 (1866) 989; Pax & K. Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 383, nom. inval. Type: Motley 480 (holotype: K), Indonesia, South Kalimantan, Bangarmassing [sic].
Twigs finely pubescent to sometimes glabrous, the hairs usually silvery short and crisped but sometimes with scattered longer erect hairs. Stipules with fine ferrugineus or silvery hairs on both surfaces particularly towards the base. Leaves: petioles densely to sparsely silvery pubescent; blades ovate, 1035 by 932 cm, deeply trilobed, sometimes with additional basal lobes or cusps, dissected to c. 2/3 or more of the leaf length, usually shallowly peltate to 13.5(4) cm, lateral lobes spreading widely and sharply, leaf lobes narrow 16 cm wide, base usually quite square and truncate to broadly rounded, margin entire, with small protruding nectaries apices acute to acuminate, adaxial surface mostly glabrous with scattered erect silvery hairs along the veins especially towards the petiole insertion, abaxial surface densely to very densely pubescent along all veins, not glaucous; 1° venation palmate with 79 prominent veins, 2° venation scalariform, looping near the leaf margins and terminating at the margin in prominent flattened nectaries; young leaves red-brown, densely covered in spiny erect red-brown hairs, soon becoming less densely hairy. Staminate inflorescences basal axes scattered with erect silvery hairs becoming glabrous, distal axes densely covered in minute ferrugineus hairs; bracts finely pubescent particularly towards base and along margins; bracteoles with both surfaces scattered to densely covered with minute red-brown hairs and yellow granular glands. Pistillate inflorescences glabrous towards the base and moderately to densely ferrugineus pubescent towards the apex; bracts ferrugineus pubescent on both surfaces, abaxial surface scattered with yellow granular glands.
Distribution Sumatra (incl. Bangka Island), Borneo (Central and South Kalimantan, Sarawak).
Habitat & Ecology A small early successional tree or large shrub occurring in primary and degraded forests from the lowlands to 1700 m.
Notes 1. Macaranga depressa forma depressa is distinguished from the other forms by the typically short pubescence on young twigs and leaves, although in some cases the twigs may be almost glabrous. The leaves of this form are often quite small, usually evenly trilobed with short angular lobes. The leaves tend to be less deeply peltate than the other two forms.
2. The one specimen from Sarawak (Clemens 20377) differs from the other collections of this form in having more strigose hairs on the abaxial leaf surface.
b. f. glabra Whitmore
Macaranga depressa (Mόll.Arg.) Mόll.Arg. f. glabra Whitmore, Kew Bull. 29 (1974) 446; S.J. Davies, Harvard Pap. Bot. 6 (2001) 398. Type: Meijer SAN 61642 (holotype: K; isotypes: L, SING), Malaysia, Sabah, Jalan Kabili, Sepilok Forest Reserve.
Twigs glabrous or rarely with scattered minute ferrugineus hairs at the nodes. Stipules glabrous, sometimes with scattered ferrugineus or silvery hairs on the base of the abaxial surface. Leaves: petioles mostly glabrous or with scattered short silvery hairs; blades ovate, 1035 by 930 cm, deeply 3- to 5-lobed, dissected to 4/5 of the leaf length, usually shallowly peltate to 14(5) cm, lateral lobes narrow and spreading widely, leaf lobes narrow 36 cm wide, base mostly truncate or very broadly rounded, margin entire, with small protruding nectaries, apices acute to acuminate, adaxial surface glabrous or rarely with scattered erect silver hairs on the veins near the petiole insertion, abaxial surface moderately to densely pubescent on the veins with short silvery hairs, not glaucous; 1° venation palmate with (7)810 prominent veins, 2° venation scalariform, looping near the leaf margins and terminating at the margin in prominent conical nectaries; young leaves densely covered in spiny erect red-brown hairs, soon becoming glabrous. Staminate inflorescences with basal axes mostly glabrous or sometimes with scattered erect silvery hairs, distal axes densely covered in minute ferrugineus hairs; bracts ovate elliptic, 514 by 47 mm, margin entire, apex acute, finely ferrugineus pubescent particularly towards base and along margins, caducous; bracteoles with both surfaces scattered to densely covered with minute red-brown hairs and yellow granular glands. Pistillate inflorescences glabrous on basal axes, with scattered minute ferrugineus hairs towards the apex.
Distribution Borneo (Sabah, Sarawak).
Habitat & Ecology A small early successional tree common in primary and secondary lowland to submontane dipterocarp forests. This form of Macaranga depressa is known from Sabah east of the Gunung Kinabalu Range, where it has been mostly collected in the lowlands. It has also been found in submontane forest up to 1300 m in the Kelabit Highlands in northern Sarawak.
Notes 1. Macaranga depressa forma glabra is distinguished from the other forms of this species by having predominantly glabrous twigs and petioles, although in some specimens the petioles have some scattered fine hairs. The leaves of this form tend to be much more deeply dissected than the other forms: the leaves frequently have five prominent lobes. The leaf margins also tend to be more strongly toothed than in the other forms.
2. The two collections from the Kelabit Highlands in Sarawak (Nooteboom 1877; Davies 99007) in addition to being collected from a higher elevation than other known collections of this forma glabra, have extremely deeply dissected leaves with long slender lobes. In other respects these specimens match the form, but further assessment of the variation within this taxon is required. It is interesting to note that the forma strigosa and forma glabra therefore appear to be sympatric in the Kelabit Highlands.
c. f. strigosa Whitmore
Macaranga depressa (Mόll.Arg.) Mόll.Arg. f. strigosa Whitmore, Kew Bull. 29 (1974) 446; S.J. Davies, Harvard Pap. Bot. 6 (2001) 399. Type: Ridley s.n. (holotype: K; isotype: SING), Malaysia, Sarawak, Matang Woods.
Macaranga divergens Mόll.Arg. in DC., Prodr. 15, 2 (1866) 989. Type: Korthals s.n. Herb. Lugd. Bat. (holotype: L; isotype: BO), Indonesia, Borneo.
Twigs densely strigose, with 23 mm long erect hyaline hairs, often also with shorter erect silvery hairs. Stipules finely pubescent and often with scattered long erect hairs. Leaves: petioles densely pubescent with long erect strigose hairs and shorter silvery hairs; blades ovate to broadly ovate, 1030(35) by 825 cm, variably trilobed, sometimes almost entire to shortly tricusped to quite deeply trilobed and sometimes with additional basal cusps or lobes, peltate to (1)26 cm, lateral lobes varying from short acute cusps to spreading narrow lobes 25 cm wide, base rounded to broadly rounded to almost truncate, margin entire, with small protruding nectaries, apices acute to acuminate, adaxial surface scattered to densely covered with erect hyaline strigose hairs particularly along the veins and towards the petiole insertion, abaxial surface densely to very densely strigose, not glaucous; 1° venation palmate with 79 prominent veins, 2° venation scalariform, looping near the leaf margins and terminating at the margin in prominent flattened and conical nectaries; young leaves mostly green or reddish-brown when very young, very densely covered in erect strigose silvery or reddish-brown hairs. Staminate inflorescences with basal axes scattered to evenly covered with erect strigose hairs, sometimes becoming glabrous, distal axes often glabrous or moderately to densely covered in minute ferrugineus hairs near flower clusters; bracts finely pubescent particularly towards base and along margins; bracteoles with both surfaces scattered to densely covered with minute red-brown hairs and yellow granular glands. Pistillate inflorescences with scattered erect strigose hairs towards the base and sometimes with minute ferrugineus hairs towards the apex; bracts ferrugineus pubescent on both surfaces, abaxial surface scattered with yellow granular glands.
Distribution Borneo (Central, East and West Kalimantan, Sarawak).
Habitat & Ecology A small early successional tree of mixed dipterocarp forests. This form of M. depressa grows in both dry areas and in areas on the margins of swamp forest. As with the other forms it has a broad altitudinal range, occurring from near sea level to c. 1400 m.
Note This form of M. depressa is perhaps the most striking and easily recognized form due to the prominent strigose hairs on the twigs, petioles, leaves and young shoots. The leaves of forma strigosa tend to be least deeply dissected of the three forms, with a rather large central leaf lamina. This is particularly so for plants at high elevation, where the plants may have almost entire leaves. The montane specimens also tend to have the most densely pubescent leaves and shoots.
Macaranga didymocarpa Whitmore, Kew Bull. 39 (1984) 607; Gen. Macaranga Prodr. (2008) 122. Type: Talib Bidin SAN 80729 (holo K; iso SING), Borneo, Sahab, Beaufort Dist., Klias.
Small trees to 9 m with white or grey bole. Twigs slender, 4 mm diam., terete, coarsely shallowly striate, finely pubescent plus a few long hairs. Stipules ovate, 4 by 2 mm, apex acute, finely velvety, papery, erect, soon caducous. Leaves: petiole 8 cm, very slender, 2 mm diam., terete, finely ochraceous pubescent plus a few long hairs; blades ovate, c. 11 by 8 cm, papery, base broadly rounded, 2 cm peltate, margin weakly crenate and undulate, apex acute, drying mid greyish brown, below slightly glaucous and granular glandular with main nerves pubescent. Staminate inflorescences amongst and behind leaves, crowded dense 6 cm furfuraceous racemes or panicles with 23 axis orders; secondary axes divaricate, to 15 mm long; peduncle 1 cm; main axis slender, 1.5 mm diam., tapering upwards, branches terminating in 7 by 5 mm catkin-like clusters of overlapping erect persistent bracts; latter ovate, 6 by 5 mm, apex acute, irregularly coarsely glandular-toothed, strongly concave, finely pubescent, papery, pinnately nerved, and enclosing a 34 mm axis with strongly overlapping bracteoles; latter oblong, 12 mm long, broadly acute, sometimes coarsely toothed, papery, pubescent, erect, persistent, longer than and enclosing a tight cluster of 34 tiny pedicellate flowers. Staminate flowers with 3 stamens, anthers 4-locular. Infructescences behind leaves, crowded, 7 cm spikes or sparsely short-branched racemes, main axis terete, ridged, ochraceous furfuraceous; peduncle 2 cm; branches alternate, 2 cm, slender; bracteoles elliptic, 3 mm, papery, ochraceous furfuraceous, mostly caducous. Fruits crowded, strongly transversely bilobed and slightly flattened, 8 by 4 by 3 mm, sessile, blackish brown, granular glandular, smooth, thinly woody; calyx tiny, unlobed, persistent; stigmas 1 mm, persistent, flattened. Seeds with sarcotesta, globose, 4 mm, testa with distant shallow pits.
Distribution Borneo (Sabah, Sarawak).
Habitat & Ecology Secondary forest, sometimes swamps.
Macaranga diepenhorstii (Miq.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 998; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 337; Whitmore, Tree Fl. Malaya 2 (1973) 111, t.8; in Airy Shaw, Kew Bull. 36 (1981) 321; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 365; Whitmore, Gen. Macaranga Prodr. (2008) 122. Mappa diepenhorstii Miq., Fl. Ind. Bat. Suppl. (1861) 457. Tanarius diepenhorstii (Miq.) Kuntze, Rev. Gen. 2 (1891) 620. Type: Diepenhorst s.n., (holo U?), Sumatra, Priaman Prov.
Small bushy crowned trees in secondary forest, or trees to 24 m tall, 70 cm diam. in primary forest. Twigs near tip drying angled, 4 mm diam.; terete further from the tip, 10(15) mm diam., smooth, glabrous, dark brown with pale stipule scars. Stipules ovate oblong, 10(15) by 5(8) mm, apex acute, cucullate, warm brown, papery, glabrous, erect, subpersistent. Leaves: petioles 16(26) cm long, 2(4) mm diam., terete, striate, furfuraceous becoming glabrous, weakly kneed at both ends; blades ovate, 1323(30) by 1018(23) cm, papery, base broadly rounded, 24(8) cm peltate, with spider's web nervation, margin crenate and distantly glandular dentate, apex acute, drying dark brown (young living leaves blackish purple), below closely finely granular glandular and glabrous except for sparse hairs on main nerves, secondary nerves curved, concolorous with lamina, tertiaries closely scalariform and clearly visible especially below. Staminate inflorescences diffuse sinuous narrow panicles to 18 cm long, 3 axis orders, glabrous; secondary axes divaricate; peduncle to 5 cm; main axis flattened, striate, 2 mm diam.; lower branches (sub)opposite, 25 cm, sinuous; bracts not seen; flower clusters on 1 cm catkin-like tertiary branches, some with a 12 mm basal stalk, towards tips these with overlapping erect persistent bracteoles subtending clusters of 48 flowers, lower bracteoles squarish, 4 by 6 mm, upper edge coarsely toothed, papery, nerved, sparsely granular glandular, strongly cucullate with at base a large dark patch (?glandular), upper bracteoles similar but smaller and entire, towards catkin base this structure repeated on 5 mm long quaternary axes. Staminate flowers globose, 0.5 mm diam.; pedicel 0.5 mm; sepals 2, glabrous; stamens 2, anthers 4-locular. Pistillate inflorescences as staminate but smaller, to 10(15) cm long and racemose, branches 23 cm, apical catkins more open; bracteoles similar; flowers solitary. Infructescences with 1 cm distant solitary fruits. Fruits dicoccous, 5 by 10 mm, the cocci sulcate, splitting, thinly woody, black granular glandular; pedicel 10 mm, glabrous; slender; sepals tiny, 0.5 mm, persistent; styles 2, 1 mm, apical, slightly flattened, recurved. Seeds round, 3 mm, coarsely ridged, with sarcotesta.
Distribution Thailand, Malay Peninsula, Sumatra (including Pulau Simeuleu and Krakatau).
Habitat & Ecology Secondary forest, also often along rivers in primary forests, occasionally on limestone, to 600m.
Note Similar to M. recurvata, see there for distinctions.
Macaranga dioica (Forster) Mόll.Arg., Linnaea 34 (1865) 199; in DC., Prodr. 15, 2 (1866) 1011; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 375; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 145; Gen. Macaranga Prodr. (2008) 124, Fig. 8. Ricinus dioica Forster, Prodr. (1786) 67 Type: Forster s.n., 1774 (iso K), Vanuatu, Tanna Island.
Mappa tannensis Spreng., Syst. Veg. 3 (1826) 878.
Macaranga urophylla Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 377; L.M.Perry, J. Arnold Arb. 34 (1953) 221.
Macaranga involucrata (Roxb.) Baill. var. glabrescens Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1012, see Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 375.
var. dioica
var. dioica: Whitmore, Gen. Macaranga Prodr. (2008) 124. Type: as the species.
Small trees to 9m. Twigs to 3 mm diam., slightly angled to almost terete, densely pale tomentose becoming almost glabrous, sometimes slightly glaucous. Stipules oblong elliptic, 10 by 34 mm, acuminate, papery, tomentose, erect, subpersistent. Leaves: petioles 713 cm, 23 mm diam., terete, tomentose; blades ovate to deltoid, 721 by 620 cm (without apex), papery, base broadly rounded to subtruncate, 5mm closed cordulate at petiole insertion, or very rarely to 1 cm peltate, with 0 to few to many small glands near insertion, margin repand with small inconspicuous glandular teeth to sometimes weakly crenate, apex caudate, 210 cm, rarely with 2 caudate lobes, drying warm brown, below velvety with simple and tufted hairs on all nerves, above slightly velvety with nerves hairy and lamina scabrid, 3 main and 24 minor nerves at petiole insertion. Staminate inflorescences narrow tomentose racemes to 11 cm; branches divaricate to 15 mm long, 12 cm apart, arising from a flower cluster; peduncle 34 cm; main axis very slightly flattened, 1.5 mm diam., basal branches subopposite; bracts tiny, elliptic, 2 by 1 mm, tomentose outside, persistent; flower clusters 2 mm, at first crowded later 5 mm spaced; bracteoles spreading, persistent, enclosing cluster, ovate, 1.5 by 1.5 mm, apex acute, cucullate, tomentose outside, glabrous inside, persistent. Staminate flowers c. 6 per cluster, sessile, granular glandular; stamens 6, anthers 4-locular. Infructescences spikes to 10 cm with apical and subapical fruit clusters; peduncle 1 mm diam., terete, tomentose, subtended by fairly persistent leafy, elliptic bracts, to 20 by 5 mm, apex acuminate, shortly stalked, pinnately nerved, tomentose, each subtending 1 fruit. Fruits globose, sometimes with a shallow suture, 5 mm, thinly woody, densely invested in 1 mm woolly soft spines and densely black glandular; pedicel 5 mm, tomentose; calyx persistent; style lateral, thread-like, to 10 mm, hirsute, plumose on one side, caducous. Seeds round, coarsely verrucose with sarcotesta.
Distribution East New Guinea, Bismarck Archipelago, Solomon Islands, Santa Cruz Group (the commonest species on Vanikoro), Vanuatu.
Habitat & Ecology Lowlands, riverbanks and secondary forest, to 210m.
Notes 1. Ricinus dioica was described from a collection (K!) made by Forster (1786) in 1774 on Tanna island, Vanuatu during Cook's second circumnavigation.
2. Close to Macaranga similis but can be distinguished by its narrower stipules and infructescence bracts and the fruits retaining their spines; the leaves are never so strongly velvety on either surface. M. galorei is also close.
3. This species is variable. It encompasses M. urophylla which Perry (1953) noted to be variable.
var. glabra Whitmore
var. glabra Whitmore, Gen. Macaranga Prodr. (2008) 126. Type: Whitmore BSIP 2227 (holo K), Solomon Islands, Santa Isabel, Maringe Lagoon, Buala Village.
Distinct in being totally glabrous except for sometimes a few hairs on the midrib below.
Distribution Papua New Guinea (New Ireland, Bougainville), Solomon Islands.
Habitat & Ecology Secondary forest. Lowlands to 950m.
Notes 1. A few var. dioica gatherings from New Guinea are almost glabrous, so I give it no higher rank.
2. Gideon LAE 77169, 77170 from 950 m on the Lelet plateau, New Ireland, are provisionally placed here. The fruits differ in possessing coarse branching spines on the upper part.
Macaranga domatiosa Airy Shaw, Kew Bull. 25 (1971) 540; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)145, Pl. 6., t.4; Gen. Macaranga Prodr. (2008) 126, Fig. 9. Type: Flenley ANU 2663 (holo K; iso L), Papua New Guinea, Western Highlands, Wabag, Ambun Valley.
Macaranga caryocarpa Airy Shaw, Kew Bull. 25 (1971) 541.
Macaranga rhodonema Airy Shaw, Kew Bull. 25 (1971) 542; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)156.
Treelets or small trees, 38(11) m tall. Twigs slender, 2vmm diam., angled, finely pubescent to subglabrous. Stipules elliptic, 5 by 2 mm, apex acuminate, papery, pubescent, erect, caducous. Leaves: petioles 2 cm, slender, 1 mm diam., terete, ridged, sparsely pubescent, weakly kneed at top; blades ovate, c. 5.5 by 3 cm, thinly leathery, base rounded, minutely cordulate, occasionally with glands near petiole insertion, margin recurved upper half with small protruding glandular teeth, apex an abrupt 1 cm caudex, drying grey-green, main nerves below sparsely granular-glandular and sparsely to densely tomentose, below sparsely granular-glandular, with 3 basal nerves, nerves and reticulations paler than lamina on both surfaces, raised, weakly bullate between the 56 pairs of secondary nerves, these looping and joining near margin and bearing (inconspicuous) axillary domatia. Staminate inflorescences delicate to 8 cm sparsely branched racemes; branches to 2 cm, divaricate, arising from a flower cluster, lowest sometimes opposite; peduncle 2 cm; main axis terete, slender, 1 mm diam., pubescent; bracteoles minute, triangular, 1 mm, acute, or 2 mm including a subulate tip, or rarely elliptic, 5 mm, papery, glabrous, persistent, smaller or longer than the 23-flowered irregularly spaced clusters. Staminate flowers reddish, developing in sequence, large, sessile; sepals 2, red-nerved, 23 mm long, acuminate; stamens 1225, anthers 4-locular. Pistillate inflorescences 515 cm delicate spikes, with 34 subulate 12 mm bracteoles and a single terminal flower. Pistillate flowers reddish, with 3.5 mm spathaceous calyx half enclosing the narrowly ellipsoid 3 by 1.5 mm ovary, narrowing into 1.5 cm style, often curved or hooked. Fruits ellipsoid, to 10 by 45 mm, leathery, sparsely granular-glandular, with persistent style, pedicel 10 mm. Seeds ovoid 7 by 5mm.
Distribution East New Guinea.
Habitat & Ecology Lower montane sometimes disturbed forest, 20603000m.
Note Distinctive in its ovate, sparsely nerved leaves which commonly bear domatia. The pistillate parts are described from Airy Shaw. The delicate sparsely flowered inflorescences place this with M. gracilis, M. kostermansii, M. misimae and M. suleensis, also small montane trees, but the big fruit with apical style is strongly different.
Macaranga ducis Whitmore, Kew Bull. 34 (1979) 602; in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 145; Gen. Macaranga Prodr. (2008) 128. Type: Brass 28890 (holo K; iso L), Papua New Guinea, Milne Bay Prov., Modewa Bay, Modewa, Gara River.
var. ducis
var. ducis: Whitmore, Gen. Macaranga Prodr. (2008) 128. Type: as the species.
Trees 525 m. Twigs 3 mm diam., angled, pubescent with mostly short and some long hairs, with conspicuous stipule scars. Stipules at base broader than twig, large, oblong-triangular, 1215 by 10 mm, acute, strongly cucullate, chestnut brown, papery, pubscent as twigs, erect, persistent. Leaves: petioles 8 cm, slender, 2 mm diam., terete, weakly pubescent; blades broadly ovate, 1317 by 811cm, thinly leathery, base broadly rounded to truncate to broadly shallowly cordate, with 1several glands near petiole insertion, margin (weakly) crenate, apex caudate, 2 cm, usually drying chestnut brown, below with sparse stiff hairs on main nerves and (sometimes sparse) black granular glands. Staminate inflorescences dense stout panicles to 14 by 7 cm, 3 axis orders, pubescent; branches divaricate; peduncle 2 cm, flattened and coarsely ridged, 2 mm diam.; branches usually alternate, rarely opposite; bracts variable triangular to elliptic, 48 mm long, 1 mm caudate, papery, pubescent, erect, subpersistent; flower clusters close with overlapping bracteoles towards branch tips; bracteoles much longer than clusters, ovate-trapezoid, 4 by 3 mm, acute, slightly cucullate, slightly fleshy, margins slightly repand, outer surface with small scattered black granular glands, erect, persistent. Staminate flowers 9 per cluster, pedicellate, 0.4 mm diam., with a few granular glands; stamens 4, anthers 4-locular. Infructescences 20 cm spikes or sparsely branched racemes, furfuraceous, axis angled, fruits in clusters of 23, subtended by caducous leafy elliptic 15 mm bracts. Fruits round, to 1 cm diam., leathery, with dense soft flat almost glabrous spines; pedicel stout 13 mm, furfuraceous; calyx persistent; style plumose. Seeds unknown.
Distribution New Guinea (Papuan peninsula, Eastern Highlands, Sepik).
Habitat & Ecology River banks and open primary rain forest to 1550m.
var. glabra Whitmore
var. glabra Whitmore, Gen. Macaranga Prodr. (2008) 129. Lelean NGF 46370 (holo K; iso L, SING), Papua New Guinea, Central Dist., Tapini subdist., above Tapini.
All parts glabrous. Stipules large, broader than twig. Leaves: blades chartaceous. Staminate inflorescences: bracteoles greatly exceeding the flower clusters. Staminate flowers: stamens 2.
Distribution Papua New Guinea (Central Prov.).
Habitat & Ecology Riverside, 900m.
Note A species distinctive in its large stipules broader than the twig and male bracteoles greatly exceeding the flower cluster. The variety is also distinctive.
Macaranga endertii Whitmore, Kew Bull. 29 (1974) 449, in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 157; Gen. Macaranga Prodr. (2008) 130. Type: Endert 5267 (holo K), Borneo, Kalimantan, West Kutai, Kombeng.
Shrubby treelets or trees to 30 m tall, 60 cm diam. Twigs solid, slender, terete, finely pubescent or becoming glabrous. Stipules narrowly triangular, 9 by 2 mm, papery, finely buff-furfuraceous, erect, caducous or subpersistent. Leaves: petioles 611 cm, slender, terete, finely pubescent or glabrous; blades broadly ovate, 816 by 512 cm, leathery, base 1 cm peltate, broadly truncate, with 1(2) glands near petiole insertion, margin distinctly crenate and with distant protruding small glands, apex acute or acuminate, 3 main and several minor nerves at petiole insertion, drying warm brown or greenish above, below densely granular glandular and main nerves sparsely or densely pubescent. Staminate not seen. Pistillate inflorescences crowded, 612 cm long, narrow open racemes with 1025 mm distant slender divaricate side branches, without axillary branches, finely pubescent or glabrous; main axis flattened; peduncle 1540 mm; bracts elliptic, 2 mm, distantly toothed, furfuraceous, caducous. Pistillate flowers in terminal umbellate cluster of c. 5 and solitary 5 mm spaced laterally, subtended by caducous bracteoles; latter linear, 23 mm long, bearing 1few large patellar glands. Fruits (? young) weakly bilobed, 3 mm diam., thinly woody, black granular glandular; pedicel thread-like, 4 mm.; calyx tiny, persistent; styles 2, caducous, recurved, 1 mm, terminal. Seeds round, with sarcotesta, surface smooth(?).
Distribution Borneo (Kalimantan; Sarawak: Miri).
Habitat & Ecology Slik: In primary forest or on exposed places. Mostly found on limestone. Up to 700 m altitude.
Note 1. This can be an unusually large tree for Macaranga.
2. Differs conspicuously from M. costulata in the very broad strongly leathery distinctly crenate leaf and more robust pistillate inflorescences.
Macaranga eymae L.M.Perry, J. Arnold Arb. 34 (1953) 200; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 145; Gen. Macaranga Prodr. (2008) 130. Type: Eyma 4966 (holo A; iso BO, K, SING), Indonesia, Papua, Wissel Lakes.
Small 8 m trees. Twigs slender, 3 mm diam., angular near tip, sparsely tomentose with long golden hairs, slightly glaucous with stipule scars. Stipules united, apparently single, forming a 7 by 1 cm gibbous apical bud, narrowing from centre to acuminate tip, densely long golden tomentose, papery, erect. Leaves: petioles 4 cm, terete, 2 mm diam., densely golden tomentose, weakly kneed at both ends; blades elliptic, 1426 by 47 cm, broadest near base, leathery, base rounded, cordulate with several glands near petiole insertion, margin finely bluntly serrate from bullations, apex long tapering to acuminate, drying chocolate brown above and glabrous except for tomentose midrib, densely golden tomentose below, strongly very finely bullate. Staminate inflorescences behind leaves, (young) narrow tomentose, 6 cm racemes; peduncle 2 cm; branches 15 cm, adpressed, alternate; bracts linear-elliptic, caducous, with crowded tiny flower clusters; bracteoles with 1 by 0.5 mm hirsute stalk and 1 by 1 mm sagittate acute head bearing c. 6 patellar glands, fleshy, spreading, persistent, smaller than cluster. Staminate flowers 67 per cluster, subsessile; stamens 6, anthers 4-locular. Infructescences slender tomentose 5 cm spikes with a single terminal fruit, axis terete, 1 mm diam.; bracts caducous, narrow to 1 cm long, with patellar glands. Fruits bilocular, 1 by 2 cm, granular glandular and very densely invested in flattish sinuous 3 mm hirsute spines; pedicel 1 cm, tomentose; calyx caducous; styles 2, linear, plumose, 6 mm, caducous.
Distribution West New Guinea (Wissel Lakes).
Habitat & Ecology 1700m.
Note Highly distinctive in its group from its very finely bullate elliptic leaves. Part of the description is taken from Perry (1953).
Macaranga fallacina Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv, add. VI (1919) 31; L.M.Perry, J. Arnold Arb. 34 (1953) 209; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)146; Gen. Macaranga Prodr. (2008) 132. Type: Ledermann 6985 (iso K), Papua New Guinea, Sepik Region, Malu.
Shrubs or small trees to 8 m. Twigs slender, 4 mm diam., slightly angular, at first rusty furfuraceous, soon glabrous, occasionally glaucous, with stipule scars. Stipules united, apparently single as an erect, elliptic, acuminate apical bud, 1222 by 5 mm, rusty pubescent, papery, caducous. Leaves: petioles 812 cm, terete, 2 mm diam., usually glabrous, occasionally furfuraceous or glaucous, kneed at both ends; blades ovate to ovate oblong, (12)21(22) by (8)13(20) cm, papery, base very broadly truncate, sometimes weakly rounded, 11.5 cm peltate with 35 glands on descending main nerves near petiole insertion, margin repand with regular distant small prominent glandular teeth, apex acute, acuminate or rarely 2 cm caudate, drying grey to pale brown, above glabrous, below (weakly) scurfy on main nerves and with sparse black granular glands, basal secondary nerves normal to midrib, curving near margin, with equally strong tertiaries descending from it to basal margin. Staminate inflorescences panicles to 20 by 5 cm, 34 axis orders, furfuraceous or glabrous; main axis flattened, 1.5 mm diam.; peduncle 13 cm; some branches opposite, without axillary branches, sometimes arising from a flower cluster, secondaries divaricate to ascending, 410 cm; flower clusters in irregular groups; bracteoles ovate oblong, 2 by 1 mm, apex acute or including a caducous spreading acumen, fleshy, glabrous, usually with adaxial patellar glands, margins sinuous, smaller than cluster. Staminate flowers c. 20, developing in sequence, sessile; stamens 6, anthers 3-locular. Infructescences spikes or open sparsely branched racemes to 18 cm long, axis slightly flattened, 2 mm diam., furfuraceous; branches opposite, ascending, to 18 cm long, bearing 45 cm distant clusters of 12 fruits; bracts stipule-like, to 15 by 3 mm, glandless, at nodes (soon caducous) and subtending fruit clusters (caducous). Fruits bilocular, 6 by 10 mm, somewhat flattened, woody, blackish, closely granular glandular and with sinuous soft flattish hairy at first dense spines; pedicel stout, 1 mm; calyx persistent; styles apical, 3 mm, plumose, soon caducous. Seeds globose, 6 mm, coarsely irregularly warty, with sarcotesta.
Distribution Throughout New Guinea.
Habitat & Ecology Low lying, sometimes swampy, primary or secondary forest, to 750 m.
Notes 1. The leaf nervation and surface glands are highly distinctive.
2. Lam 1189 has unusually large staminate panicles, with the ultimate axes thread-like and zigzag between the flower clusters.
Macaranga fragrans L.M.Perry, J. Arnold Arb. 34 (1953) 235; Airy Shaw Kew Bull.23(1969)94,95 passim; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 146; Gen. Macaranga Prodr. (2008) 133. Type: Brass 7329 (holo A; iso L), Papua New Guinea, Western Prov., Palmer River.
Big trees to 37m tall, 50 m diam. Twigs stout, 14 mm diam., roundish with conspicuous stipule and leaf scars, glabrous. Stipules broadly oblong-elliptic, to 7 cm long, 1.5 cm broad at base, tapering regularly to acuminate tip, drying mid brown, papery, variously scurfy, with a central keel, erect, persistent as a tuft protruding beyond apical bud. Leaves: petioles 47 cm, stout, 8 mm diam., rounded above except flattened in lower part, shallowly coarsely striate, glabrous, kneed at top; blades big, ovate, to 50(76) by 34(48) cm or more, papery, base broadly rounded to 15 cm peltate, margin weakly repand, apex acute, drying dark brown, completely glabrous, below with very sparse black granular glands. Staminate inflorescences open long panicles to 40 by 13 cm, scurfy at least distally; main axis flattened, striate; peduncle 24 cm long, 3 mm diam.; basal secondary axes and a few others opposite, 7 cm, divaricate; a few secondaries and tertiaries with axillary branches, tertiaries divaricate, to 3 cm; quaternaries a few, 1 cm; flower clusters at branch tips, regularly 2 mm apart at anthesis; bracteoles spreading ovate, to 10 by 5 mm, apex rounded, with nerves ending in 12 mm narrow gland-tipped teeth, scurfy on both surfaces, thick, sides somewhat incurved. Staminate flowers fragrant, 8 per cluster, globose, 0.5 mm; pedicel 0.5 mm; calyx scurfy; stamens 6, anthers 4-locular. Infructescences 3443 cm panicles, peduncle 512 cm. Fruits 2(3)-locular, tomentellose, loculi 9 mm long by 6 mm, with 0.52 mm spines mixed with minute ones; stigma 2 mm, densely plumose. Seeds almost smooth.
Distribution Throughout New Guinea (Idenburg river, Palmer river and Sepik).
Habitat & Ecology Lowland secondary forests.
Notes 1. Collected three times. The infructescence, fruit and seed descriptions are taken from Perry (1953).
2. Macaranga fragrans has a superficial resemblance to M. magnifolia from the big leaves and crowded terminal flower clusters. However, the stipules are scurfy not hirsute, the leaves are completely glabrous, the staminate bracteoles (nerved in both species) are larger at anthesis and are more strongly toothed. Moreover, the fruit ornament differs, spiny here, with curious pale warts in M. magnifolia.
3. There is a resemblance in inflorescence structure of both species to M. mappa and M. gigantea as pointed out by Airy Shaw (1969).
Macaranga galorei Whitmore, Kew Bull. 34 (1979) 602; in Airy Shaw, Kew Bull., Add. Ser. 8 (1980) 146; Gen. Macaranga Prodr. (2008) 136. Type: Craven & Schodde 1011 (holo K; iso L), Papua New Guinea, Gulf Dist., Middle Tauri River.
Shrubs, 4 m. Twigs fairly stout, 4 mm diam., terete, densely pale tomentose. Stipules elliptic, 10 mm, 2 mm wide at base, tapering to a long filiform tip, pale tomentose, erect, subpersistent. Leaves: petioles 48 cm, terete, 1.53 mm diam., densely pale tomentose; blades deltoid, broader near base 1222 by 817 cm, sometimes tricusped or shallowly lobed, papery or leathery, base broadly cuneate to truncate, cordulate, usually with a few pale glands on main nerves near base, margin repand and inconspicuously glandular dentate apex acute to acuminate, drying pale khaki, weakly velvety, all nerves below, and main nerves and surface above sparsely hairy to densely pubescent, most hairs single, a few tufted, black granular glandular below, base with 3 main and several minor nerves, weakly bullate. Staminate inflorescences racemes or 3 axis-order panicles, to 9 cm long, pale tomentose; branches divaricate, tertiaries arising from a flower cluster; peduncle c. 4cm, flattened, 1mm diam.; lowermost branches subopposite; bracts ellipcit, 2 mm, persistent, both sides hairy; flower clusters crowded; bracteoles spreading beyond cluster, ovate, 2 by 2 mm plus 1 mm blunt acumen, sometimes slightly lobed, tomentose outside, inside glabrous with 2 patellar glandse. Staminate flowers 9 per cluster, 0.8 mm, weakly pubescent and with a few granular glands; pedicel 0.2 mm; stamens 6, anthers 4-locular. Infructescences among leaves, 513 cm spikes, axis terete to slightly flattened, pubescent, with an apical and sometimes 12 subapical clusters of 36 fruits subtended by several nerved leafy pale tomentose bracts; latter deltoid, c. 15 by 15 mm, shortly petiolate, acuminate. Fruits black, globose, 5 mm diam., thinly woody, with evanescent black granular glands and short stout tomentose caducous spines; pedicel stout, 3 mm, tomentose; calyx adherent to fruit, persistent; style single, lateral, 38 mm, variably pectinate, caducous. Seeds round, verrucose, with sarcotesta.
Distribution East New Guinea (Morobe and Gulf Districts).
Habitat & Ecology Primary or secondary often alluvial lowland rain forest.
Note Close to M. similis but distinct in its elliptic stipules, leaves less velvety at least above, details of staminate inflorescence and staminate bracteoles, and the pubescent staminate flowers. Macaranga dioica is also close. The length and pectination of the style is variable.
Macaranga gigantea (Rchb.f. & Zoll.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 995; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 307; Corner, Ways. Trees Malaya (1940) 265; Whitmore, Malay. Nat. J. 20 (1967) 94, 96; Airy Shaw, Kew Bull. 26 (1972) 288; Whitmore, Tree Fl. Malaya 2 (1973) 107; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 157; Kew Bull. 36 (1981) 321; Airy Shaw, Kew Bull. 37 (1982) 26; S.J.Davies, Harvard Pap. Bot. 6 (2001) 427; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 367, Fig. 15; Whitmore, Gen. Macaranga Prodr. (2008) 137, Fig. 11. Mappa gigantea Rchb.f. & Zoll., Linnaea 29 (1857) 465. Lectotype (designated by S.J.Davies, 2001): Teijsmann s.n. (G-DC), Indonesia, Sumatra, near Muara Enim, Palembang Province.
Mappa megalophylla Mόll.Arg., Flora 47 (1864) 467. Macaranga megalophylla (Mόll.Arg.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 995; Hook.f., Fl. Brit. India 5 (1877) 449; Ridl., Fl. Malay. Penins. 3 (1924) 298. Type: Griffith KD 4724 (holotype: K, A [picture]), Malaysia, Malay Peninsula.
Mappa macrophylla Kurz ex Teijsm. & Binnend., Natuurk. Tijdschr. Ned.-Indiλ 27 (1864) 44. Lectotype (designated by S.J.Davies, 2001): Teijsmann HB 3656 (BO), Indonesia, Sumatra, Palembang.
Mappa rugosa Mόll.Arg., Linnaea 34 (1865) 197. Macaranga rugosa (Mόll.Arg.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 995. Type: Wallich 7807 (K-W, not seen), India Orientali [sic].
Macaranga incisa Gage, Rec. Bot. Surv. India 9 (1922) 245; Ridl., Fl. Malay. Penins. 3 (1924) 301; Whitmore & Airy Shaw, Kew Bull. 25 (1971) 241. Lectotype (designated by S.J.Davies, 2001): Ridley 408 (SING), Malaysia, Selangor, Kuala Lumpur.
Enormously abundant pioneer tree to 30 m tall and 3040 cm dbh, usually smaller and flowering from 510 m; twigs stout, 1025 mm in diam., terete, rugose, with elongate lenticels and conspicuous leaf and stipule scars, scattered with minute ferrugineus hairs and fine silvery grey simple or tufted hairs. Stipules huge, ovate to oblong, 1560 by 1040 mm, even larger in saplings, broadly erect, apex acute to rounded, light green, sparsely to densely covered in silvery and ferrugineus hairs, persistent in a cluster at the shoot apex, soon drying black and chartaceous. Leaves: petioles 2040(50) cm long, stout, to 9 mm in diam., strongly flattened on the adaxial surface and D-shaped in section, pubescent with coarse simple and tufted silvery hairs; blades huge, broadly ovate, 2060 by 2050 cm, often much larger in saplings, thickly coriaceous, trilobed, dissected to 1/31/2, deeply peltate to 213 cm, base broadly rounded, margin coarsely dentate, apices acute, adaxial surface sparsely to densely pubescent along the veins particularly towards the petiole insertion and the lamina scattered with minute simple ferrugineus and tufted silvery hairs, abaxial surface densely pubescent with erect simple silvery hairs and densely punctate glandular, greyish to slightly glaucous below; 1° venation palmate with 58 prominent veins, 2° venation scalariform looping and terminating at the margins in large downward-facing conical nectaries, 3°4° venation densely scalariform, 5° venation weakly scalariform, finest nerves densely reticulate and prominent; young leaves reddish-brown, densely tomentose on both surfaces. Staminate inflorescences paniculate, erect, 1550 by 1030 cm, all axes densely silvery to ferrugineus pubescent with tufted and simple hairs, the main basal axis less densely pubescent, 4 axis orders, main axis flattened, branching from less than 1/2 of the total inflorescence length, first pair of secondary branches alternate without accessory branches; bracts narrowly ovate, 410 by 25 mm, apex acute, finely silvery pubescent on both surfaces, persistent; flower clusters with 615 flowers, spirally arranged and usually in groups of 26 on the ultimate inflorescence branches; bracteoles ovate to broadly ovate, 46 by 25 mm, margin deeply irregularly dentate with 48 spreading unequal teeth, apex broadly acute, veins slightly raised on abaxial surface, both surfaces densely covered with fine silvery and ferrugineus hairs. Staminate flowers c. 1 mm long, shortly pedicellate; sepals 3, free, sparsely to densely ferrugineus pubescent; stamens 23; anthers 4-locular. Pistillate inflorescences paniculate, erect, 1225 by 612 cm, densely branched, densely silvery pubescent on all axes, 3 axis orders; bracts ovate, 35 mm long, margin entire, finely silvery pubescent, caducous. Pistillate flowers c. 2 mm long; calyx urceolate, densely minutely golden pubescent; ovary 2-carpellate; styles c. 1 mm long, free, persistent; stigma not dissected. Fruits 45 by 68 mm, bilocular, locules subglobose, pedicellate, evenly covered in yellow sticky glandular exudate, without processes, with scattered silvery hairs between the locules; pedicel 46 mm long, finely silvery to golden hairy. Seeds c. 4 mm in diam., lenticular, black, with shallow grooves, encased in a fleshy pale purple aril.
Distribution Peninsular Thailand, Malay Peninsula, Sumatra (incl. Bangka), Borneo.
Habitat & Ecology Macaranga gigantea has a very broad ecological range occurring from lowland coastal and swampy areas, to c. 8001000 m. It is one of the earliest colonists of degraded land, but can also be found in large forest gaps within primary forest. It is regularly found to be one of the dominant tree species in regenerating forests after 1020 years following the abandonment of shifting cultivation (Kochummen, Malayan Forester 29, 1966, 170181). Under good soil conditions this species can grow exceptionally quickly. It produces huge quantities of small seeds that are taken by a wide range of small birds and squirrels.
Notes 1. The only Teijsmann collection Davies (2001) has seen from Palembang is Teijsmann HB 3656 and it fits the description perfectly, and was therefore chosen as the lectotype for Mappa macrophylla.
2. Macaranga gigantea is one of the most easily recognized of all Malaysian trees. It occurs along roadsides throughout west Malesia. The enormous trilobed leaves, and thick stems with huge persistent, papery stipules make this species highly distinctive. In saplings, which are often seen along roadsides, the leaves can be as much as one meter long.
3. The flower, fruit and seed morphology of M. gigantea are characteristic of section Pruinosae. Within section Pruinosae, M. gigantea is most closely related to M. pentaloba. The differences between these two species are described under that species.
Macaranga glaberrima (Hassk.) Airy Shaw, Kew Bull. 19 (1965) 322, pro parte; Kew Bull. 23 (1969) 107; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 146; Gen. Macaranga Prodr. (2008) 137. Rottlera glaberrima Hassk., Flora 25, Beibl. 2 (1842) 41 (diagn.); Cat. Hort. Bot. Bogor. (1844) 239 (diagn.); Pl. Jav. Rar. (1848) 264 (descr.). Syntypes: Editors: No type cited, but Whitmore has listed Reichenbach f. & Zollinger 3667 and 3867.
Rottlera subfalcata Rchb.f. & Zoll., Verh. Natuurk. Vereen. Ned.-Indiλ 1 (1856) 11, 29; Linnaea 28 (1856) 311. Macaranga subfalcata (Rchb.f. & Zoll.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1007; J;J.Sm., Meded. Dept.Landbouw Ned.-Indiλ 10 (Bijdr.12) (1910) 482; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 364; Backer & Bakh.f., Fl. Java 1 (1963) 487.
Macaranga haplostachya Pax & K.Hoffm. in Engl.,Pflanzenr. IV.147.xiv add. VI (1919) 25; L.M.Perry, J. Arnold Arb. 34 (1953) 250; Airy Shaw Kew Bull. 25 (1971) 531 passim; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 162.
var. glaberrima
var. glaberrima: Whitmore, Gen. Macaranga Prodr. (2008) 137. Type: as the species.
Shrubs, treelets to 2 m or small trees to 15 m. Twigs slender, 1.53 mm diam., terete, glabrous. Stipules and bud scales subulate, 5 mm, chestnut brown, glabrous, erect, crowded at twig tip, soon caducous. Leaves: petioles 2.53 cm, slender, 1 mm diam., terete, glabrous, with a small conspicuous apical knee; blades elliptic, 810(15) by 2.54(7) cm, thickly papery, base cuneate minutely cordulate with 2 glands near petiole insertion, margin weakly sinuate, apex abruptly acuminate, 2 cm, usually drying grey green, glabrous, granular glands absent or sometimes sparse. Staminate inflorescences often as witches' brooms, spikes (but rarely sparsely once branched) to 6 cm, pubescent; peduncle to 1 cm, terete, smooth or ridged, slender; flower clusters in groups; bracteoles ovate, 1 mm, apex blunt or acute, finely toothed, persistent, strongly concave, spreading, finely pubescent outside, smaller than cluster of c. 4 flowers. Staminate flowers 1 mm, pedicellate; sepals pubescent; stamens c. 10, anthers 4-locular. Infructescences delicate 6 cm spikes, with a single terminal fruit; axis slender, 1 mm, terete, ridged, glabrous; bracts and bracteoles not seen. Fruits strongly bilobed, lobes globose, each 5 mm diam., woody, sparsely granular glandular, with a few small 1 mm spines; pedicel to 7 mm, glabrous; calyx persistent; styles not seen. Seeds globose, 5 mm, smooth, with a thin sarcotesta.
Distribution Java, Sulawesi (Palu), Nusa Tenggara (Manggarai, Timor), West New Guinea, Bismarck Archipelago (New Ireland).
Habitat & Ecology Primary forest, mostly 4501100 m.
Note Macaranga haplostachya is only known from two syntypes, Ledermann 8912, 9020. If these are ever found Airy Shaw conjectured it will prove to be M. glaberrima. I concur, despite some differences in its description from the above.
var. schoddei Airy Shaw
var. schoddei Airy Shaw, Kew Bull. 23 (1969) 107; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 146; Gen. Macaranga Prodr. (2008) 139. Type: Schodde 2884 (holo K), Papua New Guinea, Central Prov., Kokoda Trail, 5 miles N of Sogeri.
Differs solely in the short pubescence of twigs, petioles and midribs below.
Distribution Java, Nusa Tenggara (Flores), New Guinea (Papua).
Note The BO sheet of Kostermans 22118 is doubly aberrant. It has the staminate inflorescence sparsely branched, viz. racemose not spicate, and also bears fruiting spikes. Airy Shaw noted that Schodde 3054 also is hermaphrodite, a rare occurrence in Macaranga, cf. M. inamoena, M. subdentata).
Macaranga glandibracteolata S.J.Davies, Kew Bull. 54 (1999) 147; Harvard Pap. Bot. 6 (2001) 399. Type: Davies 93027 (holotype: SAN; isotype: A), Malaysia, Sabah, Danum Valley.
Tree to 25 m tall and c. 35 cm dbh, with an open spreading crown; twigs 512 mm in diam., glabrous usually with scattered ferrugineus hairs towards the apex, densely glaucous, hollow, housing ants. Bark pale-grey, smooth, hoop-marked. Stipules deltate, 68 by 57 mm, producing food-bodies on abaxial surface, 13 pairs present on shoots. Leaves: petioles terete, slender, 1825 cm long, scattered to evenly covered with minute silvery hairs particularly towards the apex; blades broadly ovate, 2545 by 1635 cm, >3 cm peltate, shallowly tricusped to trilobed, dissected to 1/101/5 of the leaf length, base very broadly rounded, margin entire, secondary veins terminating in c. 1 mm diameter protruding conical nectaries, apices broadly acute to shortly acuminate, adaxial surface glabrous, smooth, abaxial surface finely pubescent with longer silvery hairs and minute crisped ferrugineus hairs, glaucous; 1° venation palmate with 78 prominent veins, 2°4° venation prominent and densely scalariform; young leaves finely red-brown pubescent. Staminate inflorescences paniculate, erect, 2030 by 2030 cm, finely pubescent with silvery and ferrugineus hairs, the hairs more scattered to almost absent near the base, glaucous, up to 5 axis orders, first pair of secondary branches ± opposite with accessory branches; bracts deltate, 47 mm long, ± persistent, margin entire, apex acute, rarely terminated by a small conical nectary, abaxial surface densely pubescent; flower clusters of 410 flowers decussately arranged in groups of 610 at branch apices; bracteoles ovate, 36 by 34 mm, enclosing the flowers, margin finely pectinate with the central lobe elongated 12 mm with a small conical nectary at apex, apex acute, abaxial surface finely pubescent. Staminate flowers 11.5 mm long, sessile; sepals splitting to the base, apex pubescent; stamens 34; anthers 3-locular. Pistillate inflorescences paniculate, erect, 512 by 510 cm, stout, up to 3 orders of branching, secondary branches ± opposite, very finely red-brown pubescent; bracts caducous. Pistillate flowers 3.55.5 by 23 mm, solitary in bracteole axils, pedicel developing with the fruit; calyx c. 4 mm long, urceolate, densely and finely red-brown pubescent, persistent; ovary 23 by c. 2 mm, 45-carpellate, minutely pubescent at the apex and on the sutures; styles 12 mm long, fused at the base, free and spreading from 1/41/3 of the length, persisting to form a 2 mm long crown on the fruit apex; stigma not dissected. Fruits subglobose, 57 by c. 10 mm, pedicellate, one discrete glandular patch covering most of each carpel wall and developing into a uniformly raised disc covered in yellow, sticky exudate; pedicel 48 mm long, glabrous or scattered with minute ferrugineus hairs, glaucous. Seeds subtriangular-ovoid, 45 mm in diam., black, shallowly and finely grooved and pitted, encased in a fleshy aril, colour not seen.
Distribution Borneo (East Kalimantan, Sabah). The species is only known from areas in Sabah east of the Crocker Range and in East Kalimantan, it is particularly common around Danum Valley and further to the east.
Habitat & Ecology Early successional tree in lowland mixed dipterocarp forest. It grows to be quite a large pioneer tree, occurring mostly on the edges of primary forest and in logged forest. It does not appear to be very common in areas that have been farmed or severely degraded.
Note This species is most closely related to M. motleyana and M. constricta. With these species it shares glaucous twigs, 34-staminate flowers, a conical nectary at the apex of the staminate bracteole, a distinctly pedicellate fruit, and one glandular patch on each carpel wall that develops into flat, disc-like structure in the fruit. Macaranga glandibracteolata differs from these two species in having larger, more hairy and only slightly tricusped to shallowly trilobed leaves with much more prominent 2°4° venation.
Macaranga gracilis Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv add. VI (1919) 31; L.M.Perry, J. Arnold Arb. 34 (1953) 250; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 147; Gen. Macaranga Prodr. (2008) 139. Type: Ledermann 11609 (not seen), Papua New Guinea, Madang Prov., Schraderberg.
Small trees 310 m. Twigs slender, 2 mm diam., angled, glabrous. Stipules elliptic, 5 by 1 mm, apex acuminate, pale brown, papery, glabrous, erect, soon caducous. Leaves: petioles 1.5 by 3.5 cm, slender, 1 mm diam., terete, glabrous, strongly kneed at top; blades elliptic, 4.59.5 by 1.52.5 cm, thinly leathery, base gradually narrowing to rounded, with or without a few tiny scattered glands near petiole insertion, margin repand, apex caudate, 2 cm, drying grey green, glabrous, granular glands usually (very) sparse and inconspicuous, secondary nerves c. 4 pairs, distant, strongly upcurving, lowest pair at petiole insertion and extending half up blade, tertiary nerves usually ladder-like and at 90Ί to midrib, tertiaries and secondaries sometimes paler than lamina at least on underside, a few domatia sometimes present. Staminate inflorescences delicate, sparsely branched glabrous racemes to 3 cm; branches alternate, divaricate, to 1 cm, mostly arising from a flower cluster; peduncle 210 mm; main axis very slender, <1 mm terete; flower clusters irregularly to 5 mm spaced; bracteoles minute, triangular, 0.6 mm, fleshy, glabrous, spreading persistent, smaller than 13 flowered clusters. Staminate flowers developing sequentially, to 1.2 mm diam., sessile; sepals fused; stamens c.12, anther 4-locular. Infructescences as staminate, delicate 6 cm spikes or sparsely branched racemes. Fruits single or paired, terminal on branches, ellipsoid, 7 by 4 mm, densely golden pubescent, sessile; calyx persistent; style short, lateral, 1 mm. Seeds not seen.
Distribution Scattered in New Guinea.
Habitat & Ecology Primary or secondary montane forest, 15002200 m. Rare.
Note I place Sayer 12528 here. It differs in 45 flowers per staminate cluster, rusty scurfy twigs and petioles, and 1 mm peltate leaves, which dry dark brown, but otherwise fits, notably in the tertiary nerves, scalariform and at 90Ί to the midrib.
Macaranga grandifolia (Blanco) Merr., Philipp. J. Sci. 7 (1912) 394; Sp. Blancoan. (1918) 226; Interp. Herb. Amboin. (1917) 320; Enum. Philipp. Flow. Pl. (1923) 441; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 34; Whitmore, Gen. Macaranga Prodr. (2008) 142. Croton grandifolius Blanco, Fl. Filip. (1837) 753. Neotype (Whitmore, 2008): Merrill Species Blancoanae 651 (BO, K, L, W), Philippines, Luzon, Rizal Prov., Antipolo.
Macaranga porteana Andrι, Rev. Hortic. 60 (1888)176, Fig.36; Bot. Mag. 121 (1895) t. 7407; Merr., Philipp. J. Sc. Bot. 7 (1912) 238.
Macaranga mappa auct. non (L.) Mόll.Arg.: Fern.-Vill., Novis App. (1880) 195; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 320, Pl. 54A; Merr., Interp. Herb. Amboin. (1917) 29, 320; Airy Shaw, Kew Bull. 23 (1969) 94.
Trees. Twigs stout, 15 mm diam., with big prominent leaf scars, lenticellate, terete, glabrous. Stipules big, oblong-elliptic, to 8 by 3 cm, apex rounded to acute, warm brown, papery, at first with fine sericeous hairs, crowded at twig tips, long persistent. Leaves: petioles 35 cm, stout, angled, striate, finely furfurescent; blades huge, ovate, 40 by 35 cm, thinly leathery, base broadly rounded, 12 cm peltate, margin weakly distantly crenate and with small glands, apex acute, drying dull brown, below velvety when young, later glabrous except whitish hairs on nerves, with granular glands, close reticulations variously developed. Staminate inflorescences completely glabrous permanently dense panicles on a basal 25 cm bare peduncle, to 10 cm, eventually expanding to 20 cm, 3 branching orders, axes flattened; main axis bracts persistent, conspicuous, papery, elliptic, to 4 by 1 cm, apex acuminate, entire, smaller upwards; flower clusters close, some contiguous; bracteoles persistent, concave, ovate, 3 mm, apex acute, entire, papery. Staminate flowers 48 per cluster, shortly pedicellate, 0.8 mm diam.; sepals free;, stamens 23(5), anthers 4-locular. Infructescences spherical axillary, sessile masses of fruits, 6 cm across. Fruits with 2 strong ovoid lobes, 6 mm long, woody, drying blackish or warm brown, with apically branching tentacular processes to 1cm long, process bases and wall closely granular glandular; pedicel 1cm; calyx small, persistent; styles 1/3 fused, apical, 5 mm. Seeds spherical, coarsely pitted.
Distribution Philippines (Luzon, Mindoro). Naturalised on Hawaii (Hilo, Honolulu).
Notes 1. Very distinctive when fertile. There are two Philippines' Macaranga with gigantic leaves. Sterile, M. grandifolia of Luzon and Mindoro is very similar to M. mappa of Luzon, Samar and Mindanao, but has somewhat smaller leaves without tufted hairs below. Inflorescences and fruits are quite different.
2. Merrill (1917, 1918) has described how Mόller (1866: 1000) mistakenly equated Croton grandifolius Blanco of Luzon to a similar big-leaved species Folium mappae Rumph. of Ambon, and he created the combination M. grandifolia for the Philippines' species.
Macaranga griffithiana Mόll.Arg. in DC., Prodr. 15, 2 (1866) 993; Hook. f., Fl. Brit. India 5 (1887) 452; Pax & K. Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 309; Ridl., Fl. Malay. Penins. 3 (1924) 299; Corner, Ways. Trees Malaya (194) 265; Whitmore, Malay. Nat. Journ. 20 (1967) 97; Tree Fl. Malaya 2 (1973) 147;Airy Shaw, Kew Bull. 26 (1972) 289; S.J. Davies, Harvard Pap. Bot. 6 (2001) 400; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 367, Fig. 16. Macaranga motleyana (Mόll.Arg.) Mόll.Arg. subsp. griffithiana (Mόll.Arg.) Whitmore, Kew Bull. 29 (1974) 448. Type: Griftith KD 4713 (holotype: K; isotype: A [scrap]), Malaysia, Melaka.
Macaranga adenophila Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 310. Type: Johs. Schmidt 280 (holotype: C, not seen), Thailand, Koh Shan Island (= Koh Chang).
Slender glaucous tree to 15(20) m tall with branches clustered towards the ends of the axes, in open areas often shrubby and flowering from 45 m tall; twigs 49 mm in diam., glabrous or rarely with a few scattered minute ferrugineus hairs towards the apex, densely glaucous, hollow, housing ants, usually exuding some reddish clear latex when cut. Bark smooth, hoop marked, mottled pale grey-brown and white. Stipules rather small, ovate, 46 by 67 mm, the pair not completely encircling the twig, recurved, succulent, producing food-bodies on the abaxial surface, 14 pairs present on the shoots but often caducous. Leaves: petioles terete, slender, 615(23) cm long, glaucous, glabrous; blades ovate, 1025(28) by (8)1023 cm, usually shallowly trilobed, dissected to 1/51/3 of the leaf length, central lobe broadly deltate c. 48(10) cm wide at the base, lateral lobes 13(6) cm wide short and spreading with shallow rounded sinuses, 25 cm peltate, base narrowly rounded, margin entire, apices shortly acuminate, adaxial surface glabrous, abaxial surface glabrous to finely pubescent on the veins, densely glaucous; 1° venation palmate with 67(8) prominent veins, 2° venation scalariform looping near the leaf margins and terminating at the margin in tiny protruding conical nectaries, 2°4° venation only feintly visible not arising prominently from the abaxial leaf surface; young leaves red-brown, minutely ferrugineus pubescent on the adaxial surface and densely yellow granular glandular on the abaxial surface when young, soon becoming glabrous. Staminate inflorescences paniculate, erect, 1025 by 1018 cm, basal axes mostly glabrous and densely glaucous, distal axes sparsely to densely ferrugineus pubescent, up to 4 axis orders, main unbranched axis 28 cm long flattened, first pair of secondary branches ± opposite with accessory branches; bracts elliptic, 715 by 37 mm, usually persistent beyond anthesis, bright yellow-green when fresh, drying black, margin entire, apex acute, scattered with minute ferrugineus hairs towards the base, otherwise glabrous; flower clusters with c. 10 flowers, decussately arranged and strongly grouped on the ultimate inflorescence branches; bracteoles ovate to broadly ovate, 35 by 2.54.0 mm, enclosing flower clusters, margin erose to very shallowly pectinate, apex acute and usually terminated by a narrowly caudate apex c. 1 mm long with a tiny conical nectary, both surfaces scattered to densely covered with minute silvery to red-brown hairs and yellow granular glands, adaxial surface also with a dense patch of minute ferrugineus hairs at the base. Staminate flowers c. 1 mm long, shortly pedicellate; sepals free, splitting irregularly to the base at anthesis, apex usually scattered with minute red-brown hairs; stamens 23; anthers 4-locular. Pistillate inflorescences paniculate, erect, 416 by 38 cm, stout, basal axes glabrous and densely glaucous, distal axes covered in fine ferrugineus hairs, 23 axis orders, secondary branches ± opposite without accessory branches; bracts caducous, not seen. Pistillate flowers c. 34 mm long, solitary in bract axils; calyx urceolate, 23 mm long, densely covered with fine ferrugineus and silvery hairs, persistent, splitting irregularly as ovary expands; ovary 4(5)-carpellate, c. 2 mm long; styles c. 2 mm long, fused at the base, free and spreading from c. 1/3 of the length, persisting to form a prominent 23 mm long crown at the fruit apex; stigma not dissected. Fruits subglobose, compressed, 79 by 812 mm, pedicellate, sutures bluish and glaucous, one discrete glandular patch on each carpel wall developing into a slightly raised flat disc covered in yellow, sticky exudate; pedicel 24 mm long, scattered with minute ferrugineus hairs, glaucous. Seeds subtriangular-ovoid, 3.54 mm in diam., black, with shallow coarse grooves and a small cruciform scar, encased in a fleshy bright red aril.
Distribution Laos, S Vietnam, Thailand (Southeast and Peninsular), Malay Peninsula, Sumatra.
Habitat & Ecology An early successional tree common in degraded areas, which also grows into a tall slender tree along rivers in primary forest. It is most common as a bushy treelet or shrub in swampy secondary forest. It is often found in freshwater swampy areas, but may also be seen on the margins of peat swamp forest. The species is very light demanding and is often seen in wet areas in heavily degraded forest.
Notes 1. Macaranga griffithiana is closely related to M. motleyana and M. constricta. These species share glaucous twigs, staminate bracteoles terminated by one small conical nectary, and 45-carpellate fruits with one disc-shaped glandular patch per locule. Macaranga constricta differs from the other two species in having much longer and erect inflorescences, and entire acuminate staminate bracteoles. Whitmore (1974) considered M. griffithiana a subspecies of M. motleyana. These species differ in a range of morphological and ecological characteristics. The leaves of M. griffithiana are shallowly dissected with short spreading cusps and broad sinuses, whereas in M. motleyana the leaves are long and ascending with deep narrow sinuses. The styles of M. griffithiana are very shortly connate and spreading from about 1/3 of the length, whereas they are connate to almost 2/3 of the length in M. motleyana. The bracteoles of the staminate inflorescences are only shortly and irregularly toothed in M. griffithiana whereas they are deeply and evenly pectinate in M. motleyana. The distributions of the two species do not overlap: M. motleyana is restricted to Borneo and M. griffithiana occurs from Indochina, south through the Thai-Malay peninsula to Sumatra.
2. Davies (2001) was not been able to find the type of M. adenophila. From the description and the location he suspects this is in fact M. griffithiana, the only species of section Pachystemon known from southeast Thailand.
Macaranga hartleyana Whitmore, Gen. Macaranga Prodr. (2008) 144. Type: Hartley 10694 (holo K), Papua New Guinea, Central Prov., Western Section, Kakoda Road at Owens Corner.
Small trees to 10 m. Twigs slender, 3 mm diam., terete, striate, sparsely pubescent. Stipules elliptic, 5 by 1.5 mm, densely finely pubescent, erect, soon caducous. Leaves: petioles 24 cm, slender, 1 mm diam., terete, striate, densely finely pubescent, weakly kneed at both ends; blades elliptic, 6.512 by 24 cm, thinly leathery, base cuneate minutely cordulate with several small glands near petiole insertion, margin repand to weakly crenate, apex tapering to a 2 cm caudex, drying dark chocolate brown or pale khaki-green, above sparsely hairy or glabrous, below with spreading hairs on lower midrib and with diffuse to almost absent black granular glands, secondary nerves c. 7 pairs, scalariform tertiaries parallel to them. Staminate inflorescences crowded and intermingling, delicate, 6 cm panicles, branching 13 mm from base into 2 divaricate narrow racemose parts, each with a few 6 mm branches; main axes angled, striate, slender 0.8 mm diam., densely finely pubescent; bracts tiny, triangular, 1 mm, pubescent, persistent; flower clusters small, regularly 7 mm spaced with 7 flowers; bracteoles ovate, 1 by 1.5 mm, apex rounded, papery, glabrous, spreading, persistent, smaller than cluster. Staminate flowers sessile, granular glandular; stamens 7, anthers 4-locular. Infructescences densely crowded and intermingled, delicate 4 cm panicles, similar to staminate. Fruits solitary, globose, 2.5 mm diam., thinly woody, densely black granular glandular; pedicel ultimately 8 mm. slender, sparsely pubescent; calyx persistent (initially enclosing), densely tomentose; style basal, 5 mm, coarsely plumose. Seeds globose, coarsely verrucose, with sarcotesta.
Habitat East New Guinea.
Habitat & Ecology Primary montane forests, 6601000m.
Macaranga havilandii Airy Shaw, Kew Bull. 23 (1969) 112; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 147; S.J.Davies, Harvard Pap. Bot. 6 (2001) 402. Type: Haviland 978 (holotype: K; isotype: SAR), Malaysia, Sarawak, near Kuching.
Macaranga havilandii Airy Shaw var. resecta Airy Shaw, Kew Bull. 25 (1971) 534. Type: Morshidi S 24096 (kolotype: K; isotypes: A, L, SAR, SING), Malaysia, Sarawak, Lambir Hills National Park.
Slender glaucous treelet to 35(6) m tall and 2.5 cm dbh with no or few secondary branches and leaves clustered at the apex of the stem; twigs 58 mm in diam., glabrous, usually densely glaucous, hollow, housing ants. Stipules ovate, 36 by 4.06.5 mm, not completely encircling the twig, recurved, succulent, producing food-bodies on abaxial surface, 26 pairs present on the shoot. Leaves: petioles terete, very long and slender, 1240 cm long, glabrous; blades ovate, 2035 by 1322 cm, shape very variable, usually deeply trilobed or 5-lobed with broad basal cusps or rarely sub-entire, usually dissected to 1/23/4 of the leaf length, deeply peltate to 510 cm, base rounded to ± truncate, margin entire or sometimes broadly sinuate, apices finely acuminate, adaxial surface dark green, glabrous, abaxial surface glabrous with minute ferrugineus hairs on veins particularly near petiole insertion, glaucous; 1° venation palmate with 67 prominent veins, 2° venation scalariform and terminating at the leaf margin in small protruding conical nectaries, 3°4° venation loosely reticulate not prominent; young leaves red-brown with minute ferrugineus hairs, soon becoming glabrous. Staminate inflorescences narrowly paniculate, erect, cruciform, 712 by 24 cm, 3 axis orders, main axis unbranched for c. 4/5 of length, first pair of secondary branches ± opposite with accessory branches, main basal axis mostly glabrous, 2° and 3° axes minutely ferrugineus pubescent; bracts ovate, 812 by 46 mm, margin entire, apex acute, glabrous or with scattered minute hairs, caducous; flower clusters with c. 10 flowers, spirally arranged and crowded at the inflorescence apex; bracteoles ovate, 57 by 23 mm, enclosing flower clusters, margin entire, apex caudate, scattered with minute crisped ferrugineus hairs. Staminate flowers c. 1 mm long, sessile; sepals fused, splitting to c. 1/3, apex with scattered minute ferrugineus hairs, stamens 1; anthers 3-locular. Pistillate inflorescences paniculate, cruciform, erect, 716 by 28 cm, 2 axis orders with one pair of opposite secondary branches, basal unbranched axis long with the flowers clustered in the distal 1/5 of inflorescence, main axis often with ant holes, glabrous at base, ferrugineus pubescent towards apex and on the branches; bracts ovate, c. 7 by 6 mm, apex acute, ± glabrous, glaucous, caducous. Pistillate flowers 46 mm long, solitary in bracteole axils; calyx urceolate, 1.52 mm long, with scattered fine ferrugineus hairs, persistent; ovary 45-carpellate, c. 1.5 mm long; styles 35 mm long, fused for 4/5 of length, free and spreading at apex, persisting to form a prominent 35 mm long crown at fruit apex; stigma not dissected. Fruits subglobose, 56 by 811 mm, bright red, shortly pedicellate, one discrete glandular patch on each carpel wall remaining smooth and not becoming horn-like, covered in pale pinkish, sticky exudate; pedicels 37 mm long, slender, glaucous, glabrous or with fine scattered ferrugineus hairs. Seeds 44.5 mm in diam., subtriangular-ovoid, black, with shallow irregular grooves and a small cruciform scar, encased in a fleshy red aril.
Distribution Borneo (Brunei, Sarawak).
Habitat & Ecology This species occurs in primary mixed dipterocarp forest and rarely in degraded forests. It has only been found on relatively nutrient-poor sandy humult ultisol soils usually in association with small streams and/or small landslides. The species is moderately high-light demanding, however it is considerably more shade-tolerant and slow growing than the other Bornean pioneer species of Macaranga. The growth form of M. havilandii is very unusual for Bornean Macaranga. It often occurs in small clumps of several slender stems of only 35 m tall arising from a single individual. The stems are fragile and appear to fall over or get pushed over by falling branches, following which the stems produce adventitious roots and several resprouts forming small clumps of stems. Individuals begin to reproduce when the stems are only one centimeter in diameter.
Notes 1. The species is readily distinguished from other Bornean Macaranga by its irregularly dissected and very delicate leaves that have a velvety dark green sheen on the upper surface. The cruciform, simply branched inflorescences with the flowers clustered at the ends is diagnostic for this species. Red fruits without raised appendages do not occur in any other Bornean Macaranga species. The red-fruited M. kingii and M. caladiifolia have two prominent ridges or horns on each carpel wall.
2. Macaranga caladiifolia is also a small treelet, however it differs from M. havilandii in having raised ridges on the fruit, flowers with 35 stamens, 4-locular anthers, stems that are irregularly swollen, and is not glaucous.
Macaranga hengkyana Whitmore, Gen. Macaranga Prodr. (2008) 146. Type: De Vogel 9564 (holo L; iso BO, K, MAN), Indonesia, Papua, Sorong, Hutan Lindung, km 14 on road to Aimas.
Small trees 7 m. Twigs squarish, stout, 5 mm diam, slightly rusty furfuraceous, glaucous, grey. Stipules elliptic, 12 by 3 mm, densely rusty tomentose, leathery, erect, caducous. Leaves: petioles to 21 cm, very slightly flat near base, 3 mm diam., distinctly glaucous, glabrous, kneed at both ends; blades big, oblong, to 38 by 26 cm, thinly leathery, base broadly truncate, minutely 2 mm peltate, with small glands between petiole insertion and margin, margin repand with a few small broad glandular teeth, apex acute, drying pale grey above, khaki green below, glabrous, below with close, lamina-coloured granular glands, nervation pinnate. Staminate inflorescesces and flowers unknown. Infructescences behind leaves, narrow distant 5 cm panicles with 2 main axes from the base, each bearing a few regularly spaced spreading branches to 15 mm long; main axes angular, 1.5 mm diam., weakly tomentose. Fruits (young) crowded at branch tips, ellipsoid or (sometimes deeply) bilobed, to 3 by 5 mm, thinly woody, densely golden granular glandular; calyx persistent, sessile, subtended by a somewhat persistent bracteole; latter elliptic, 5 by 2 mm, pubescent, leathery, spreading, sublaciniate in its upper part; style apical, short, plumose, caducous. Seeds globose, 4 mm, more or less smooth, with sarcotesta.
Distribution West New Guinea (known from the type only).
Habitat & Ecology Lowland primary forest, 15m. [Apparently swamp, see note sub M. palustris - Eds.]
Notes 1. The only species this resembles is M. palustris of western Papua, the other lowland member of the Brunneo-floccosa group, which also has a strictly pinnate leaf nervation. It is, however, amply distinct. Both are reminiscent of the Conifera species group of the Sunda Shelf.
2. Named for Hengky Turot, local expert on the Ayawasi flora.
Macaranga herculis Whitmore, Kew Bull. 39 (1984) 608; Gen. Macaranga Prodr. (2008) 147. Type: Vinas & Lelean LAE 59322 (holo K; iso L), Papua New Guinea, West Sepik Dist., near Busilmin.
Big trees, 29 m tall, 21 cm diam., clear bole 21 m. Twigs 7 mm diam., strongly angled, with stipule scars, densely pale pubescent. Stipules ovate oblong to 12(15) by 4(7) mm, abruptly narrowing to an acuminate tip, blackish brown, papery, both sides densely tomentose except glabrous at margins, erect to reflexed, caducous. Leaves: petioles 10 cm, 3 mm diam., slightly flattened above, densely pale pubescent, weakly kneed at both ends; blades ovate, 24 by 16 cm, thinly leathery, base broadly rounded, 5 cm peltate, margins repand, apex acute to abruptly broadly 1 cm acuminate, drying mid khaki brown, above with crisped hairs on main nerves, below densely velvety with crowded white short spreading hairs on all nerves, with dense small golden granular glands, spider's web nervation, tertiaries and quaternaries scalariform. Staminate inflorescences and flowers unknown. Infructescences 6 cm spikes; peduncle 4 cm; axis stout, slightly flattened, 2 mm diam., densely pale pubescent. Fruits crowded towards tip, solitary, weakly bilobed, 9 by 14 mm, thinly woody, blackish, rather sparsely granular glandular with slender sinuous 8 mm slightly flattened processes swollen into 1 mm elongate apical gland, pedicel 10(15) mm long, stout, 1 mm diam., pubescent; calyx persistent. Seeds unknown.
Distribution East New Guinea (known from the type only).
Habitat & Ecology Secondary montane rain forest, 1500 m.
Notes 1. One of the tallest of all Macaranga species, could have timber plantation potential.
2. Similar to M. tentaculata of lowland Papua, but with smaller, tomentose stipules and the fruit processes coarser, and to M. clavata, see there.
Macaranga heynei I.M.Johnson, Contrib. Gray Herb. n.s. 68 (Aug. 1923) 90; Whitmore, Biol. J. Linn. Soc. 1 (1969) 226, t. 2; Tree Fl. Malaya 2 (1973) 112; in Airy Shaw, Kew Bull. 36 (1981) 320; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 369; Whitmore, Gen. Macaranga Prodr. (2008) 149. [Rottlera montana Heyne ex Baill., Ιtude Euphorb. (1858) 430, nomen nudum]. Macaranga javanica (Blume) Mόll.Arg. var. montana Heyne ex Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1005. Macaranga montana (Heyne ex Mόll.Arg.) Heyne ex Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 321, non Merr., Philipp. J. Soc. Bot. 7 (1912) 394. Type Wallich Cat. 272 no. 7835 (holo K-WALL; iso K, P, W), Singapore.
Macaranga robiginosa Ridl., Kew Bull. (Dec. 1923) 367; Fl. Malay. Penins. 3 (1924) 303.
Macaranga javanica auct. non (Blume) Mόll.Arg.: Hook.f., Fl. Brit. India 5 (1887) 451; Corner, Ways. Trees Malaya (1940) 266.
Treelets or trees to 10m. Twigs slender, leafy part to 3 mm diam., terete, blackish, weakly rusty furfuraceous, distal inflorescence bearing parts to 1 cm diam., innovations rusty pubescent. Stipules elliptic, 10 mm long including long narrow tip, base 1.5 mm wide, rusty pubescent, erect, soon caducous. Leaves: petioles 58 cm, slender, 1 mm diam., terete, finely ridged, weakly rusty furfuraceous or glabrous, weakly kneed; blades trullate, 1018 by 4.58 cm, thickly papery, base usually truncate, occasionally broadly cuneate, rarely broadly rounded, cordulate or 1 mm peltate and with several glands at petiole insertion, margin entire minutely recurved, apex acuminate, abruptly so for 2 cm, below slightly glaucous and sometimes rusty furfuraceous on main nerves; secondary nerves strictly pinnate but basal pair arising at petiole insertion. Staminate inflorescences behind leaves, crowded sparsely branched racemes or panicles, to 14 by 6 cm, 2 or 3 axis orders, finely rusty furfurescent, branches divaricate, peduncle short 2 cm or less; main axis fairly slender, 2 mm, terete, ridged, lower branches alternate or rarely subopposite, sometimes with short axillary branches; bracts not seen, ultimate branches slender; flower clusters crowded or irregularly spaced, smaller than the subtending bracteoles; latter erect, persistent, ovate, to 10 by 6 mm, tip caudate to 2 mm, with small apical gland, distant long linear blunt gland-tipped marginal teeth, fairly thick, densely furfuraceous. Staminate flowers in clusters of c. 20, 0.2 mm diam., sessile; sepals 2, glabrous; stamens 2, anthers 4-locular. Pistillate inflorescences as staminate, but always racemes and mostly shorter, c. 6 cm, branches alternate and accessories not seen. Fruits in groups of 3 or 4, tiny, bilocular, 3 by 4 mm, close evanescent rusty granular glands; pedicel slender, 5 mm; calyx small, persistent; stigmas apical, tiny, persistent. Seeds coarsely shallowly rugose, with sarcotesta.
Distribution Peninsular Thailand, Malay Peninsula (except north east where it is replaced by M. laciniata), Sumatra (including Pulau Simeuleu).
Habitat & Ecology Open places, often gregarious, sometimes on exposed subsoil. Lowlands to 1000 m.
Note Three nerves arise from the petiole insertion but are quite clearly the midrib and the basal pair of secondaries.
Macaranga hispida (Blume) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 990; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 366, t. 61AE; in Engl., Pflanzenr. IV.147.xiv, add. VI (1919) 29; Merr., Enum. Philipp. Flow. Pl. (1923) 442; Airy Shaw, Kew Bull. 37 (1982) 26; Alph. Enum. Philipp. Is. (1983) 34; Whitmore, Gen. Macaranga Prodr. (2008) 150, Fig. 12. Mappa hispida Blume, Bijdr. (1825) 624. Rottlera hispida (Blume) Kurz ex Teijsm. & Binn., Natuurk. Tijdschr. Ned.-Indiλ 24 (1864) 45. Tanarius hispidus (Blume) Kuntze, Revis. Gen. Pl. 2 (1891) 619. Type: not indicated (L?), Moluccas.
Halecus rugosa Rumph., Herb. Amboin. 3 (1743) 198; see Merr., Interp. Herb. Amboin. (1917) 320, nom. inval.
Macaranga platyclada Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv, add. VI (1919) 30; L.M.Perry, J. Arnold Arb. 34 (1953),208; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 152.
Macaranga roemeri Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xvii, add. VII (1924) 185.
Small trees, 410 m tall with irritant hairs. Twigs tetragonal or flattish near tip, stout, 47 mm across, finely warm brown hispid, rarely (sub)glabrous. Stipules united, apparently single, erect, leathery, big and prominent, elliptic, 59 cm long, finely golden to brown hispid, amplexicaul, soon caducous, leaving a prominent scar. Leaves: petioles to 11 cm, round, fairly stout, 2 mm diam., hispid; blades ovate, sometimes slightly asymmetric, 153(40) by 720(26) cm (rarely 3-lobed), thinly leathery, base cordate with a very deep and broad sinus to 4 by 5 cm, and broadly rounded basal lobes, sometimes with a few large but inconspicuous glands at petiole insertion, margin usually with distant fine glandular teeth, apex acute sometimes to 4 cm caudate, drying mid brown; both surfaces softly velvety, below from dense spreading short mostly clustered white hairs on all nerves, above from close caducous white hairs borne singly at apex of a tiny papilla, below with sparse blackish granular glands, secondary nerves pinnate the basal pair with strong tertiaries extending into basal lobes. Staminate inflorescences finely velvety panicles to 24 cm long; peduncle 715 cm; main axis stout, flattened, 2 mm diam., tapering, branching only in distal half, sometimes with axillary branches, 3 axis orders; 3rd order slender sinuous with close 2 mm clusters of numerous tiny flowers; bracts subpersistent, conspicuous, spreading, linear-elliptic to 8 mm, larger but similar to the 2 mm caducous hispid bracteoles; latter with 1 mm stalk, widening slightly to elliptic apical half with obtuse apex, bearing 36 patellar glands in 2 rows. Staminate flowers 6 or more per cluster, 0.5 mm, shortly pedicellate; stamens 7, exserted, anthers 3-locular. Pistillate flowers: sepals elliptic, 4 mm, hispid; ovary hispid, twin 20 mm apical, papillose, caducous styles. Infructescences stout hispid spikes, 814(26) cm long, axis flattened, 3 mm wide; peduncle 715 cm; bracts ovate elliptic, 15 by 38 mm, apex acute, with base narrow, hispid on centre line and with numerous patellar glands in 2 rows, soon caducous; bracteoles similar, but smaller, to 10 mm. Fruits 1-few crowded towards tip, sometimes bilobed, 1015 mm diam., leathery, shaggy with long fine tapering hirsute soft processes, surface granular glandular; pedicel stout 23 mm; calyx caducous. Seeds lenticular, coarsely verrucose.
Distribution Philippines (throughout), Sulawasi, Moluccas and New Guinea.
Habitat & Ecology Oten riverine. Very common west of New Guinea. Lowlands to 1700 m.
Notes 1. Highly distinctive. Note the clustered hairs on the leaf below and the closely pustular upper leaf surface. Mόller (1866) mistook bracts for bracteoles, which are more evanescent, much smaller, but with the same morphology.
2. The type of M. platyclada (Ledermann 9508, K!) has staminate characters well within the range of M. hispida and the tetragonal twigs are well developed. This reduction extends the range of M. hispida into New Guinea.
3. Macaranga barkeriana, M. racemohispida and M. suwo are similar.
Macaranga hoffmannii L.M.Perry, J. Arnold Arb. 34 (1953) 242; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 162; Gen. Macaranga Prodr. (2008) 153. Syntypes: Ledermann 12685, 12957 (not seen), Papua New Guinea, Sepik area, Felsspitze.
Macaranga acuminata Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv, add. VI (1919) 28 (non Ridl., 1916 which is M. densiflora Warb.).
Trees, 1020 m tall. Twigs slender, dark, glabrous and sparsely granular-glandular. Stipules not seen, most certainly small or minute. Leaves: petioles 23.5 cm, slender, glabrous; blades ovate or ovate-elliptic, 712 by 2.54 cm, leathery, base narrowed and narrowly peltate, 3-nerved base plus 24 pairs of secondary nerves, margin entire, apex caudate acuminate, shining, glabrous above, very sparsely pilose on midrib below and very densely granular-glandular. Staminate inflorescences 46 cm panicles, divaricately branched almost at base, glabrous, granular-glandular, branches slender of more or less equal length; clusters 1020 flowered; lower bracteoles oblong, 2 mm long, upper triangular, denticulate. Staminate flowers 2 mm broadl sepals 3, elliptic, acute, outside densely granular-glandular; stamens 1012, anthers 4-locular. Pistillate unknown.
Distribution Northeast New Guinea (known from the syntypes only).
Habitat & Ecology Montane forests, 14001500 m.
Note Not seen by me. The description is paraphrased from Perry's translation, amended against the original Latin. This keys out to the Dioica group.
Macaranga hosei King ex Hook.f., Fl. Brit. India 5 (1887) 449; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 309; Ridl, Fl. Malay. Penins. 3 (1924) 298; Airy Shaw, Kew Bull. 26 (1972) 289; Whitmore, Tree Fl. Malaya 2 (1973) 111; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 152; S.J.Davies, Harvard Pap. Bot. 6 (2001) 429; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 370. Lectotype (designated by S.J.Davies, 2001): King's Collector 3599 (holo: K, A [photograph]; iso: L), Malaysia, Perak.
Macaranga pseudopruinosa Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 308. Type: Haviland & Hose 3210 (holotype: K; isotypes: L, SAR), Malaysia, Sarawak.
Large tree with a bluish-green appearance, to 2030 m tall and 3040 cm dbh, although flowering somewhat smaller; twigs 611 mm in diam., terete, glabrous, usually densely glaucous. Bark pale grey, white or light brown, prominently hoop-marked, smooth or minutely fissured. Stipules 1015 by 1015 mm, orbicular surrounding the stem, erect and perpendicular to the stem, many pairs persistent, coriaceous, not succulent, glabrous, producing food bodies on the adaxial surface. Leaves: petioles terete, slender, 1025 cm long, slightly glaucous, glabrous; blades broader than long to almost rounded, not flat, slightly convex, 820 by 1025 cm, deeply trilobed, dissected to c. 1/23/4, the lateral lobes almost the same size as the central lobe, the lateral lobes slightly asymmetrical, not peltate or rarely 15 mm peltate, base very broadly rounded, strongly cordate, margin ±entire with small conical nectaries along the basal margin and usually along all margins, the distal margin sometimes appearing wavy to dentate, apices acute or shortly acuminate, 1° venation palmate with 5 prominent veins and 23 less prominent veins arising from the petiole insertion, 2°3° veins densely scalariform, 4° venation loosely scalariform, minor veins usually prominent and finely reticulate; adaxial surface glabrous, abaxial surface with dense to rarely sparse erect silvery hairs along all veins and densely punctate glandular; young leaves green, adaxial surface glabrous or with a few minute ferrugineus hairs at the petiole insertion, abaxial surface densely covered in silvery erect sharp hairs. Staminate inflorescences erect narrow panicles often extending above the leaves, 2040 by 1020 cm, to 4 axis orders, basal unbranched axis strongly flattened and usually more than 1/2 of the total inflorescence length, first pair of secondary branches alternate without accessory branches, basal axes glabrous, distal axes with silvery to ferrugineus pubescence; bracts ovate, 57 mm long, margin entire, glabrous, often persistent; flower clusters with 615 flowers, spirally arranged on inflorescence branches; bracteoles broadly ovate, 46 by 34 mm, margin irregularly deeply dentate with spreading narrow unequal teeth, apex broadly acute, both surfaces covered in minute silvery to ferrugineus hairs. Staminate flowers c. 0.7 mm long, yellow-green, shortly pedicellate; sepals 3, free, densely pubescent; stamens 23; anthers 4-locular. Pistillate inflorescences erect narrow panicles often extending above the leaves, 1535 cm long, up to 3 axis orders, basal unbranched axis strongly flattened and usually more than 1/2 of the total inflorescence length, basal axes glabrous, distal axes with silvery to ferrugineus pubescence; bracts ovate, 510 mm long, margin entire, glabrous, often persistent. Pistillate flowers c. 1.5 mm long; calyx urceolate, c. 1 mm long, with fine silvery or ferrugineus hairs; ovary 2-carpellate; styles c. 1 mm long, free, persistent. Fruits bilocular, 45 by 56 mm, green or yellowish-green at maturity, locules subglobose, without processes, covered in yellowish-green, sticky exudate; pedicel slender, 58 mm long, with fine silvery or ferrugineus hairs. Seeds c. 3 mm in diam., lenticular, black, shallowly pitted, encased in a fleshy violet aril.
Distribution Peninsular Thailand, Malay Peninsula, Sumatra, Borneo (yet to be collected in Sabah, where it appears to be replaced there by the ecologically similar and closely-related M. pearsonii. S.J.Davies (2001) expects that M. hosei might be found in southwest Sabah, outside the range of M. pearsonii).
Habitat & Ecology Macaranga hosei is a common early successional tree in lowland forests throughout west Malesia. Macaranga hosei occurs on a wide range of substrates from nutrient-poor sandy soils to richer shale-derived soils, although it seems to be more abundant on the former particularly in Borneo. It can also be found on soils across a wide range of soil water levels. In some areas it occupies the margins of swampy ground, although not usually in swamps, and in other areas it occurs on raised ridges of apparently low soil water status. Macaranga hosei is a very high-light demanding pioneer tree and grows quickly in good conditions. It is one of a group of species that grows to be a relatively large tree. It can be found in degraded land, logged forest and in large openings within primary forest where it may persist for approximately 30 years.
Notes 1. Macaranga hosei is a core member of section Pruinosae and shares with other species in the section bicarpellate fruits, and irregularly dentate staminate bracteoles, along with several other features.
2. Macaranga hosei is most closely related to M. rufescens. These species share evenly trilobed apeltate leaves, orbicular glabrous persistent stipules that are perpendicularly erect from the stem, and glabrous main inflorescence axes. Macaranga hosei differs from M. rufescens in having much longer inflorescences which have along basal unbranched rhachis, dense erect spiny hairs along the veins on the abaxial leaf surfaces, more densely glaucous twigs, smaller leaves with the central lobe often appearing almost dentate towards the apex, prominent conical nectaries along the base and apex of the leaf margin, and glabrous upper surfaces to young leaves. In the herbarium these two species may be confused at first, but in the field there is no doubt of their distinctness. Macaranga hosei has a bluish-green appearance to the canopy and all leaves are dark green. In contrast, the canopy of M. rufescens has a reddish-green appearance, with the younger leaves often being ferrugineus and the petioles red. In several parts of Sarawak the two species grow in sympatry; M. hosei is often found on slightly poorer sites than M. rufescens. There is no evidence of hybridization between these species.
Macaranga hullettii King ex Hook.f., Fl. Brit. Ind. 5 (1887) 452; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 383; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 147. S.J.Davies, Harvard Pap. Bot. 6 (2001) 404; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 370. Type: King's Collector 2907 (holotype: CAL, not seen; isotypes: K, L), Malaysia, Perak.
Macaranga bartlettii Merr., Papers Michigan Acad. Sci. 19 (1934) 161. Type: Bartlett 7656 (holotype: L; isotype: NY), Indonesia, Sumatra, east coast, near Aek Sordang, Koealoe.
Macaranga hullettii King ex Hook.f. subsp. borneensis Whitmore, Kew Bull. 29 (1974) 446. Type: Haviland 3229 (holotype: K; isotype: SAR), Malaysia, Sarawak, near Kuching.
Macaranga cornuta auct. non Mόll.Arg.; Corner, Ways. Trees Malaya (1940) 300.
Small tree to 18 m tall, but often flowering from as small as 3 m; twigs 59 mm in diam., usually glabrous or with scattered to dense erect silvery hairs, hollow, housing ants. Bark smooth, pale grey-brown, hoop-marked. Stipules broadly ovate-elliptic, 59 by 510 mm, mostly glabrous or with scattered minute silvery or ferrugineus hairs near the base, light green or yellowish-brown when fresh rarely darker, recurved and appressed to the stem, the margin usually slightly upturned in dry specimens, producing food-bodies on the abaxial surface, usually c. 34 pairs persistent on the stem. Leaves: petioles terete, 820(25) cm long, mostly glabrous or sometimes finely silvery pubescent; blades narrowly ovate to ovate, 1432 by 616(22) cm, unlobed or tricusped to sometimes trilobed, the lobes usually short and narrowly acuminate, or rarely dissected to c. 1/3 of the leaf length with ascending sharp lobes, 13(4) cm peltate, rounded to broadly rounded, margin entire to distantly and prominently serrate with small flattened nectaries, apex acute to finely acuminate, adaxial surface glabrous or scattered with erect silvery hairs along the veins and particularly toward the petiole insertion, abaxial surface usually glabrous or sometimes with fine silvery hairs along the veins, 1° venation palmate with 79 prominent veins, 2° venation scalariform and looping near the margins or terminating at the margin in small protruding nectaries, 3° venation scalariform, 4° venation loosely reticulate; young leaves pale to reddish brown, with fine ferrugineus hairs and sometimes with scattered erect silvery hairs, soon becoming glabrous. Staminate inflorescences paniculate, erect, 1030 cm long, light green drying black, glabrous rarely with a few minute ferrugineus hairs towards the apex, 3(4) axis orders, basal unbranched axis flattened and 1/23/4 of total inflorescence length, first pair of branches opposite with accessory branches; bracts ovate-elliptic, 613 by 36 min, margin entire, apex acute, glabrous or scattered with fine ferrugineus hairs towards the base, caducous; bracteoles ovate-elliptic, 48 by 35 mm, margin entire or often with several lateral narrow teeth, apex caudate, adaxial surface often densely ferrugineus scaly particularly towards the base, abaxial surface glabrous or scattered with minute ferrugineus hairs. Staminate flowers 2035 per cluster, c. 1 mm long, sessile; sepals 3, fused, splitting to c. 1/4, apex densely covered in minute crisped ferrugineus hairs; stamens 1, anthers 3-locular. Pistillate inflorescences 410 by 25 cm, unbranched with the flowers/fruits clustered at the distal end of the axis, or sometimes with one pair of short opposite branches without accessory branches, usually glabrous or with erect silvery hairs along the basal axis; bracts finely dissected, caducous. Pistillate flowers 35 by c. 2 mm; calyx urceolate, c. 2 mm long, ± glabrous or with a few scattered ferrugineus hairs, persistent: ovary (4)5-carpellate, c. 2 mm long; styles 23 mm long, expanding at maturity, fused at the base, free and spreading from c. 2/3 of the length, persisting to form a prominent 35 mm long crown at fruit apex. Fruits subglobose, 68 by 1013 mm, sessile, one discrete glandular patch on each carpel wall developing into a long slender horn-like process 47 mm long, covered in greenish-yellow, sticky exudate. Seeds ovoid, 44.5 mm in diam., black, coarsely grooved, encased in a fleshy bright pinkish-red aril.
Distribution Peninsular Thailand, Malay Peninsula, Sumatra, Borneo (Brunei, Kalimantan, Sarawak).
Habitat & Ecology A widespread early successional species occurring in primary and secondary lowland forests, and up to 1350 m in submontane forests. This species is most commonly found between 400 and 1000 m along the edges of streams, often in valleys and steep habitats, in dryland forests. It occurs on both sandy and clay-rich soils, although it appears to have a preference for soils over sandy substrates. This species is not one of the very high-light demanding species of Macaranga; it has medium growth and photosynthetic rates, and is often found in partial shade within primary mixed dipterocarp forests.
Notes 1. Macaranga hullettii is distinguished from other core members of section Pachystemon by the combination of several features: entire or only shallowly dissected narrowly ovate leaves, fruits with very long horn-like processes when mature, long erect staminate inflorescences, and long caudate apices to the staminate bracteoles often with multiple shallow or deep lobes. It appears to be closely related to Macaranga triloba from which it differs in leaf shape, in the morphology of the staminate bracteole, in the lack of densely strigose hairs on the leaf bud, and in having hollow ant-inhabited stems.
2. Whitmore (1974) recognized two subspecies of M. hullettii, one from Malaya and Sumatra, and one from Borneo, based on the degree of dissection of the staminate bracteoles. The Bornean material often has very deeply dissected bracteoles with multiple lobes, and the Malay specimens often have a single caudate apex with only shallow lateral teeth, but this character varies significantly within areas and multiple-lobed bracteoles occur in Malaya, Sumatra and Borneo. Bracteoles with sub-entire margins also occur in Borneo. On the basis of the more extensive collections now available, it is clear that these subspecies are not distinct.
3. There is however considerable morphological variation within M. hullettii as it is currently recognized. The degree of pubescence, leaf dissection, and staminate inflorescence morphology all vary considerably within this species. Plants with quite deeply lobed leaves appear to be common in Johore in the southern Malayan peninsula and in parts of Sumatra, but less common elsewhere in the Peninsula and in Borneo. In Sumatra the dissected leaves look quite similar to those of M. triloba. In Borneo there is also considerable morphological variation within M. hullettii. The Bornean specimens are often finely pubescent on the petioles and the lower leaf surfaces, although again there are glabrous specimens from Borneo and pubescent specimens in Sumatra and the Malay peninsula. Davies (2001) suspects that detailed analysis of these patterns may reveal some gene flow between M. hullettii and other closely related taxa, particularly within Sumatra. In addition in Borneo, Davies (2001) suspects that some of the specimens may represent new species. Further field work, particularly in submontane areas in Borneo, is required.
Macaranga hypoleuca (Rchb.f. & Zoll.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 992; Hook.f., Fl. Brit. India 5 (1887) 448; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 311; Ridl., Fl. Malay. Penins. 3 (1924) 3004; Corner, Ways. Trees Malaya (1940) 266; Airy Shaw, Kew Bull. 26 (1972) 289; Whitmore, Tree Fl. Malaya 2 (1973): 106; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 147; S.J.Davies, Harvard Pap. Bot. 6 (2001) 407; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 371. Mappa ? hypoleuca Rchb.f. & Zoll., Acta Soc. Regiae Sci. Indo-Neerl. 1 (1856) 30; Linnaea 28 (1857) 309. Type: Teijsmann s.n. (holotype: not seen), Indonesia, Sumatra, near Lubualang, Padang Province.
Tree to 30 m tall and 3040 cm dbh, flowering somewhat smaller; twigs 712(14) mm in diam., slightly angular at the apex especially when dry, glabrous or with scattered minute ferrugineus hairs towards the apex, intensely glaucous white. Bark bluish-green white and glaucous when young to pale brown, prominently hoop-marked, quite smooth. Stipules deltate, c. 1 by 0.7 cm, thickly coriaceous, spreading perpendicularly to the stem, glabrous or with scattered minute ferrugineus hairs at the base, glaucous, persistent or sometimes caducous. Leaves: petioles terete, slender, 1030 cm long, glabrous, glaucous; blades broader than long to almost rounded, usually slightly convex, 1024 by 1230 cm, deeply trilobed to c. 3/4 of length, rarely with additional basal cusps, the three lobes subequal with the central lobe only slightly longer and broader than the laterals, deeply peltate to 1.54 cm, base broadly rounded sometimes slightly cordate, margin entire sometimes with tiny protruding conical nectaries, apices acute or shortly acuminate, adaxial surface glabrous, abaxial surface white and intensely glaucous with scattered ferrugineus hairs on the veins; 1° venation palmate with 810 prominent veins, 2° venation strongly looping and sometimes ending in prominent conical nectaries at the leaf margin, 3° veins scalariform, 4° and finer veins indistinct due to white wax and not or only weakly scalariform; young leaves dark red-rown, densely covered in minute ferrugineus hairs and scattered with long silvery hairs, soon becoming glabrous, producing food-bodies on lower surface. Staminate inflorescences erect panicles, to 1028(35) cm long, up to 4 axis orders, branching from <1/2 axis length, main axis flattened, first pair of secondary branches ± opposite with accessory branches, basal axes glabrous densely glaucous, distal axes moderately to densely covered in crisped ferrugineus hairs; bracts deltate, 36 mm long, margin entire, both surfaces scattered with minute ferrugineus hairs; bracteoles ovate, 2.54 by c. 3 mm, enclosing the flower clusters, margin finely evenly pectinate with >10 short appressed teeth, apex acute, both surfaces densely covered in minute ferrugineus hairs. Staminate flowers 1015 per cluster, c. 1 mm long, shortly pedicellate; sepals 3, fused, densely covered in minute ferrugineus hairs; stamens 1; anthers 4-locular. Pistillate inflorescences 515 cm long, 2 axis orders, branched from near the base, branches short, densely glaucous, basal axes glabrous to scattered with minute ferrugineus hairs, distal axes densely ferrugineus pubescent; bracts ovate-elliptic, 57 mm long, entire or with a finely pectinate apex, covered in ferrugineus hairs, caducous. Pistillate flowers c. 23 mm long; calyx urceolate, c. 1 mm long, glaucous with minute ferrugineus hairs, ovary 3-carpellate; styles c. 1mm long, free, persistent. Fruits 3-locular, sub-globose, 56 by 68 mm, bluish-green, glaucous, two discrete glandular patches on each carpel wall developing into slightly raised disc-like ridges, covered in bright yellow, sticky exudate; pedicel 311 mm long, glaucous, with scattered minute crisped ferrugineus hairs. Seeds 34 mm in diam., spheroidal to slightly flattened, black, with large round shallow pits, encased in a fleshy bright red aril.
Distribution Peninsular Thailand, Malay Peninsula, Sumatra, Borneo.
Habitat & Ecology Macaranga hypoleuca is a common and widespread early successional tree. It grows in a wide range of habitats from the lowlands to around 1000 m. It occurs mostly in dryland dipterocarp forests but may also be seen on the edges of swampy forests. It appears to be quite tolerant of a range of soil conditions and can be found on richer shale-derived soils as well as on sandstone-derived soils. While this species flowers at relatively small sizes, it grows to over 30 m in height and is one of the largest pioneer trees in section Pachystemon. It is very high-light demanding and colonizes forests after selective logging, and regenerates abundantly following shifting agriculture.
Notes 1. Macaranga hypoleuca is one of the most distinctive species of south-east Asian Macaranga, characterized by peltate, deeply trilobed leaves which are very densely white glaucous on the lower surface. It is most closely related to M. beccariana with which it shares three carpellate fruits with each carpel bearing two waxy ridges at maturity, unistaminate flowers, erect triangular stipules, finely pectinate staminate bracteoles, and food-bodies produced on the lower surface of young leaves. It differs from M. beccariana principally in the shape of the leaf, with M. hypoleuca having broadly rounded leaves with three ovate to broadly ovate subequal lobes, whereas M. beccariana has more deeply dissected leaves with three linear to narrowly lanceolate subequal lobes.
2. Macaranga hypoleuca is superficially similar and sometimes confused with M. pruinosa. However, that species is restricted to peat swamp forest, has upwardly turned rounded stipules, bicarpellate fruits, two to three stamens per flower, and the leaves are never as densely glaucous as in M. hypoleuca.
Macaranga hystrichogyne Airy Shaw, Kew Bull. 33 (1978) 68; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 147; Gen. Macaranga Prodr. (2008) 155. Type: Koster BW 13820 (holo K), Indonesia, Papua, Vogelkop, Arfak Mountains, Minjambau.
Small trees, 46 m. Twigs 3 mm diam. near tip, terete, densely persistently pale brown tomentose. Stipules triangular, 3 by 37 by 4 mm, apex acute, leathery, tomentose outside, erect, sometimes persistent. Leaves: petioles c. 8 cm, 3 mm diam., terete, pale brown tomentose, not kneed; blades triangular ovate, 1420 by 1420 cm, leathery, base truncate, 5 mm cordate with overlapping, slightly convex, often multigland-bearing lobes, margin slightly sinuous with small inconspicuous protruding glands, apex acute, drying mid or dark brown, becoming minutely bullate (appearing shagreened), above nerves densely tomentose, lamina with sparse hairs, below tomentose and black granular glandular, with 3 main and several minor basal nerves from petiole insertion. Staminate inflorescences tomentose racemes to 14 cm, branches divaricate, sinuous to 6 cm; peduncle 3 cm; main axis stout, 3 mm diam., terete; branches alternate, some arising from a flower cluster, a few with short 5 mm axillary branches; bracts triangular, 3 mm, hirsute, some persistent; flower clusters contiguous or irregularly spaced; bracteoles triangular, 2 by 2 mm, apex acute, hirsute outside, glabrous inside sometimes with several patellar glands, spreading, smaller than cluster, persistent. Staminate flowers 20 per cluster, hirsute, shortly pedicellate; stamens 610, anthers 4-locular. Infructescences 11 cm tomentose panicles with 3 branch orders; peduncle 1 cm; main axis stout, 3 mm diam., terete; lower main branches opposite, no axillary branches, secondaries 3 cm, tertiaries 1 cm; bracteoles oblong, 3 by 1 mm, apex obtuse, very thick, adaxially wholly covered by c. 6 patellar glands, abaxially hirsute, patent, persistent. Immature Fruits solitary or paired, irregularly c. 3 mm spaced, globose, 3 mm diam., black granular glandular and with dense short slender spines, body and spines sparsely hirsute; pedicel stout, hirsute, to 3 mm; calyx persistent; style single, lateral, 24 mm, curved, plumose. Seeds flattened (? immature), 2 mm diam., smooth, with sarcotesta.
Distribution New Guinea (Arfak mountains and Central Prov. of Papua New Guinea).
Habitat & Ecology Secondary or disturbed forest, 4501230 m.
Note The pistillate from west New Guinea (Koster BW 13820) provided the type specimen. Airy Shaw (1978: 69) tentatively placed the staminate (Isles & Vinas NGF 34457) here. It is a close match on leaf, twigs and indumentum, but has the larger stipules and these are more persistent. A perhaps more substantial difference is in the bracteoles. I still concur with my 1980 opinion that these two sheets are conspecific, given the polymorphism of other species of this general affinity but, like Airy Shaw, I do now have some slight doubts.
Macaranga indica Wight, Icon. Pl. Ind. Orient. 5 (1852) 23, t. 1883; Icon. Pl. Ind. Orient. 6 (1853) 5, t.1949, ii; Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1009; Kurz, Forest Fl. Brit. Burma 2 (1877) 387; Hook.f., Fl. Brit. India 5 (1887) 446; Trimen, Handbook Fl. Ceylon 4 (1898) 70; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 349, t. 58; Parkinson, Forest Fl. Andaman Is. (1923) 238; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 444; Huras. & Tanaka, Fl. E. Himalaya (1966) 179; Airy Shaw, Kew Bull. 23 (1969) 93; Whitmore & Airy Shaw, Kew Bull. 25 (1971) 241; Whitmore, Kew Bull. 25 (1971) 241; Airy Shaw, Kew Bull. 26 (1972) 290, Whitmore, Tree Fl. Malaya 2 (1973) 107; Gard. Bull. Singapore 31 (1978) 54; Chiu, Guihaia 2 (1982) 148,in key; Long, Fl. Bhutan 1 (1987) 805; Chakrab. & N.P.Balakr., J. Econ. Taxon. Bot. Addit. Ser. 9 (1992) 80; Philcox, Revised Handbook Fl. Ceylon 11 (1997) 169; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 371; Whitmore, Gen. Macaranga Prodr. (2008) 156. Possible syntypes: Wight KD 2632, Nov.1851 (K, P); Wight KD 2633 (K, P, W); both India, Neilgherries, Courtallum.
? Trewia hernandiifolia Roth, Nov. Pl. Sp. (1821) 374, cf. Hook.f., Fl. Brit. India 5 (1887) 424.
Macaranga adenantha Gagnep., Bull. Soc. Bot. France 69 (1923) 701; in Lecomte, Fl. Indo-Chine 5 (1926) 443, t. 52, 711, e descr. et tab.; Chiu, Guihaia 2 (1982) 268.
Macaranga aff. perakensis auct. non Hook.f.: Corner, Ways. Trees Malaya (1940) 268.
Small or medium trees, to 20 m. Twigs to 10 mm diam., usually terete, glabrous or furfurescent, sometimes glaucous. Stipules triangular, 8(10) by 4(6) mm, apex acute, blackish brown, papery, furfuraceous, sometimes recurved, soon caducous. Leaves: petioles 618 cm, slender, to 3 mm diam., terete, smooth, glabrous, not kneed; blades ovate, 1022 by 920 cm, papery, base broadly rounded, strongly (1.5)4 cm peltate, nearly always with a few conspicuous elongate glands on main nerves somewhat distant from petiole insertion, margin entire, apex acute to broadly acuminate, drying grey-brown, grey-green or chocolate brown, occasionally glaucous and glabrous, but usually finely pubescent when young and becoming glabrous except often for a few hairs on main nerves, sometimes remaining velvety below with matted white hairs a few to mostly tufted, secondary nerves palmate, numerous arising from petiole insertion. Staminate inflorescences amongst and behind leaves, overlapping, usually dense, racemes or occasionally panicles, 612 cm long, usually 2 occasionally 3 axis orders; peduncle 24 cm; axes (strongly) flattened, to 2 mm wide, glabrous or furfuraceous, striate; primary branches 13 cm apart, divaricate, to 3 cm long, lowest alternate, subopposite or rarely opposite, sometimes dehisced, leaving scars on peduncle; bracts not seen; branches usually strongly zigzag at the distant flower clusters, these subtended by conspicuous spreading, spoon-like caducous glabrous bracteoles with flattened 12 mm stalk and a big conspicuous subapical round 23 mm diam. patellar gland, occasionally twin glands; flower clusters often on a 12 mm elongate capitulum, this developing as a next order axis, so secondary axes (except lowest few) arising from a cluster of flowers or pedicel scars and the rare 3rd order axes arising from flower clusters. Staminate flowers in clusters of 58, 0.5 mm diam., shortly pedicellate; sepals free, glabrous except often with a few subapical granular glands; stamens 57, anthers 4-locular. Pistillate inflorescences as staminate but branches not zigzag and only 2 axis orders. Fruits 1(3) per cluster, round, to 4 mm, thinly woody, black granular-glandular, becoming smooth; pedicel 510 mm; calyx persistent; short style thread-like, eccentric caducous, sometimes a few or most fruits deeply bilobed with twin styles attached subapically near point of union. Seeds shallowly coarsely verrucose, with sarcotesta.
Distribution Sri Lanka, India (Sikkim, west Peninsular, Andaman and Nicobar Islands; see Chakrabarty & Balakrishnan, 1992), Nepal, Bhutan, south China, north and east Thailand, Laos, Viernam, Malay Peninsula (main Range south to Gunung Nuang), Sumatra (Kerinci).
Habitat & Ecology Lowlands to mountains at 2100 m.
Notes 1. A variable species. The staminate inflorescence structure and the bizarre spoon-like bracteoles are highly distinctive. Northern collections usually almost glabrous and with glaucous twigs and sometimes leaves, southern forms with the leaves velvety below, sometimes (nearly always in Malaya) with bilocular fruits and sometimes with delicate inflorescences. These variants are not discrete and do not warrant taxonomic status.
2. From Gagnepain's (1922, 1926) description M. adenantha is a northern form of M. indica.
3. If a specimen of Trewia hernandiifolia can ever be traced and it is indeed this taxon then it is a much earlier name.
4. For a note on type specimens see under M. peltata (Whitmore 2008).
Macaranga indistincta Whitmore, Kew Bull. 29 (1974) 447; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 148; S.J.Davies, Harvard Pap. Bot. 6 (2001) 409. Type: Cuadra 2274 (holotype: K; isotypes: A, SING), Malaysia, Sabah, Sandakan, Sapagaya.
Small tree to 15 m tall with an overall rather dull green appearance; twigs 59 mm in diam., densely pubescent with fine erect silver-grey hairs, hollow, housing ants. Bark pale brown, smooth. Stipules ovate, 69 by 810 mm, dark brownish to purple when fresh, recurved, succulent, adaxial surface finely pubescent, producing food-bodies on abaxial surface, 26 pairs present on shoots. Leaves: petioles terete, 1022 cm long, densely pubescent; blades ovate to broadly ovate, 1532 by 1425 cm, grey-green, shallowly tricusped to trilobed or sometimes with an additional pair of cusps at the leaf base, dissected to 1/10 to 1/5 of the leaf length or rarely more, lateral lobes narrowly acute to acuminate and spreading, central lobe broadly acute, >23 cm peltate, base broadly rounded,; leaf margin entire, apices acute, adaxial surface mostly glabrous or with scattered minute ferrugineus hairs along the veins, abaxial surface densely pubescent, hairs erect silvery, not glaucous; 1° venation palmate with 79 prominent veins, 2° venation scalariform running into the leaf margins and protruding as narrow erect conical nectaries, 3° and 4° venation densely scalariform; young leaves reddish-brown, densely pubescent. Staminate inflorescences paniculate, erect, 927 by 618 cm, scattered to evenly covered in long erect silvery hairs on the basal axis, distal axes almost glabrous, drying dark blackish brown, up to 4 axis orders, main axis flattened, first pair of secondary branches opposite with accessory branches; bracts ovate-elliptic, 47 by 34 mm, margin entire, apex acute to acuminate, glabrous, caducous; flower clusters with 1525 flowers, spirally arranged and evenly spaced along inflorescence branches; bracteoles broadly ovate, 23.5 by 2.54 mm, cupulate and enclosing flower clusters, glabrous, margin pectinate with 612 short narrow and well-spaced teeth, the sinuses twice as broad as the teeth, usually the central tooth more elongated than laterals, scattered with yellow granular glands, apex rounded, adaxial surface with a dense patch of minute ferrugineus hairs at the base. Staminate flowers c. 0.75 mm long, sessile; sepals fused, splitting irregularly to c. 1/4, apex densely covered in minute red-brown hairs; stamens 1; anthers 3-locular. Pistillate inflorescences paniculate, erect, 615(20) by 510 cm, stout, mostly glabrous distally but with scattered erect sharp-pointed silvery hairs towards base, up to 3 axis orders, secondary branches ± opposite with accessory branches; bracts caducous, not seen. Pistillate flowers 34 mm long, solitary in bracteole axils; calyx urceolate, 22.5 mm long, apex truncate, glabrous or with very few scattered fine red-brown hairs and yellow granular glands sometimes slightly glaucous, persistent, splitting irregularly as ovary expands; ovary 4-carpellate, 23 mm long, with fine hairs on the sutures; styles c. 2 mm long, fused for 4/5 of the length, free and spreading at the apex, persisting to form a prominent 23 mm long crown at the fruit apex; stigma not dissected. Fruits subglobose, 46 by 710 mm, sessile, one discrete glandular patch on each carpel wall developing into a long slender horn-like process 47 mm long, covered in yellowish green, sticky exudate. Seeds 3.54 mm in diam., subtriangular-ovoid, black, with shallow coarse grooves and a small cruciform scar, encased in a fleshy bright pink aril.
Distribution Borneo (East Kalimantan, Sabah). It is only known from areas east of the mountains of Gunung Kinabalu and the Crocker Range. One collection from near Kutai in East Kalimantan (Endert 3266) may be M. indistincta, although it is not a very good specimen and may be the closely related M. velutina.
Habitat & Ecology Macaranga indistincta is a small early successional tree locally common in secondary forest in Sabah. It occurs in the lowlands and rarely up to 1000 m. The species occurs in canopy openings within primary forest, in secondary forests and along roadsides.
Notes 1. This species is a typical ant-inhabited member of the triloba-clade within section Pachystemon. The combination of densely pubescent twigs and abaxial leaf surfaces, pectinate staminate bracteoles and very long horns on the mature fruits separates this species from all others.
2. It appears to be most closely related to M. velutina which differs in having entire staminate bracteoles and shortly horned fruits. Macaranga indistincta also has much longer staminate and pistillate inflorescences than M. velutina.
Macaranga induta L.M.Perry, J. Arnold Arb. 34 (1953) 244; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)147; Gen. Macaranga Prodr. (2008) 158. Type: Clemens 6568 (holo A), Papua New Guinea, Morobe Prov., Yunzaing.
subsp. induta
subsp. induta: Whitmore, Gen. Macaranga Prodr. (2008) 158. Type: as the species.
Trees 1017 m. Twigs 3 mm, squarish, densely pubescent to furfuraceous, becoming subglabrous. Stipules oblong linear, 36 mm, apex acute, very soon caducous. Leaves: petioles 25 cm, 1.5 mm diam., terete, striate, at first finely pubescent, kneed at both ends; blades elliptic to ovate-elliptic, 717 by 37cm, broadest near base, leathery, base cuneate to rounded, 24 mm peltate, with scattered inconspicuous glands near petiole insertion, lowest part of margin often recurved, narrowing towards 1 cm acuminate apex, drying grey above buff below, below with a fine dense whitish felted tomentum on lamina, the main nerves tomentose, close dark granular glandular, above glabrous closely finely conspicuously reticulate and sometimes minutely slightly bullate, below scalariform tertiaries and reticulum raised and conspicuous, secondary nerves 814 pairs, basal secondaries and several minor nerves arising from petiole insertion. Staminate inflorescences open pyramidal racemes, to 10 by 6 cm, axes sparsely pubescent; branches divaricate arising from a flower cluster, lower branches 3 cm; peduncle 2 cm; main axis slender, 1 mm diam., terete, striate; flower clusters contiguous or in small 3 mm evenly spaced groups; bracteoles broadly triangular, 1.5 by 2 mm, with short abrupt acumen, strongly cucullate, glabrous, papery, nerved, spreading, smaller than cluster, persistent. Staminate flowers 67 per cluster, shortly pedicellate; calyx granular glandular; stamens 12, anthers 4-locular. Pistillate inflorescences 6 cm spikes, or scarcely branched racemes with 1 cm branches arising from flower cluster, axes as staminate. Pistillate flowers 24 per cluster; clusters 2 cm distant, only 1 flower maturing. Fruits depressed globose, 6 mm across, very shallowly bilobed, leathery, densely whitish pubescent and granular glandular, sometimes with short spines (? caducous); pedicel to 11 mm, slender, sparsely granular glandular; calyx persistent; styles 2, erect, 6 mm, termina,l caducous, papillose. Seeds round, coarsely verrucose, with sarcotesta.
Distribution New Guinea (throughout), Bismarck Archipelago (New Britain).
Habitat & Ecology Primary or secondary montane forests, 13002300 m.
Note Distinctive in the whitish felt on the leaves below, which does not obscure the nervation. Stipule description from Perry (1953). The type subspecies seen only with spikes in the pistillate. Perry (1953) described short spines on the immature fruit of the type.
subsp. paucinervis Airy Shaw
subsp. paucinervis Airy Shaw, Kew Bull. 33 (1978) 70; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)147; Gen. Macaranga Prodr. (2008) 159. Type: Carr 15831 (holo K; iso G, SING), Papua New Guinea, Northern Prov., Lala River.
Differs in tree 420 m. Blades very soon glabrous below and only 58 pairs of secondary nerves, basal pair running nearly 1/3 up lamina, secondaries and reticulations only weakly raised below.
Distribution East New Guinea.
Habitat & Ecology Primary (once in disturbed) montane forest, 14002600 m.
Macaranga inermis Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 333; L.M.Perry, J. Arnold Arb. 34 (1953) 252; Airy Shaw, Kew Bull. 31 (1976) 395; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 147; Gen. Macaranga Prodr. (2008) 159. Syntypes: Schlechter 17740 (K, P), Schlechter 17902 (K, P), both Papua New Guinea, Madang prov., Kani Mountains.
Macaranga effusa Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv, add. VI (1919) 26; L.M.Perry, J. Arnold Arb. 34 (1953) 254.
Macaranga mallotiformis Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv, add. VI (1919) 27. Macaranga inermis Pax & K.Hoffm. var. mallotiformis (Pax & K.Hoffm.) L.M.Perry, J. Arnold Arb. 34 (1953) 252.
Macaranga penninervia Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv, add. VI (1919) 26. Macaranga inermis Pax & K.Hoffm. var. penninervia (Pax & K.Hoffm.) L.M.Perry, J. Arnold Arb. 34 (1953) 253.
Macaranga multiflora C.T.White, Proc. Roy. Soc. Queensland 55 (1944) 83; Airy Shaw, Kew Bull. 31 (1976) 395.
Macaranga inermis Pax & K.Hoffm. var. plurifoveata L.M.Perry, J. Arnold Arb. 34 (1953) 254.
Trees (2)15(30) m. Twigs 34 mm diam., angled, sometimes ridged, often dark brown, sometimes furfuraceous. Stipules elliptic, to 10 by 5 mm, acute, dark brown, papery, furfurescent, soon glabrous, erect, subpersistent. Leaves: petioles 315 cm long, 1.53 mm diam., terete, glabrous or weakly furfuraceous, kneed at both ends but more strongly at top; blades ovate, less often elliptic, (10)1520(26) by (4)914(19) cm, leathery, base (broadly) rounded to subtruncate, minutely cordate to 2 mm peltate, margin repand to subcrenate often, slightly recurved at least near base, often with up to 4 small close glands near petiole insertion, apex (abruptly) usually broadly acuminate, glabrous, with close small lamina-coloured granular glands. Staminate inflorescences crowded narrow racemes to 8 cm long, very weakly furfuraceous; branches to 2 cm ascending; peduncle 12 cm; main axis slender, 1 mm diam., terete, ridged, some branches opposite; bracts ovate, 1 mm, papery, persistent; flower clusters single or double, evenly spaced; bracteoles ovate, 1 by 2 mm, apex rounded, glabrous, spreading, persistent, much smaller than the flower clusters; latter 3 mm diam. 11-flowered. Staminate flowers globose, 1 mm, developing sequentially, sessile; stamens 5 becoming exserted, anthers 4-locular. Pistillate inflorescences as staminate but racemose or paniculate with 3 axis orders, secondaries 5 cm, tertiaries 1.5 cm and thread-like; flowers in evenly spaced clusters of 14, 3 mm apart; bracteoles ovate, 1 mm, apex acute, spreading, persistent. Pistillate flowers developing 3(5) mm pedicel. Fruits globose, small, 2(4) mm diam., very thinly woody, closely black granular glandular; pedicel slender, 38 mm; calyx persistent; style basal, papillose, caducous, 38 mm. Seeds globose, with shallow depressions, with sacrotesta.
Distribution New Guinea (throughout), Bismarck Archipelago (New Britain), Australia (Queensland).
Habitat & Ecology Mostly primary forest, mostly mountains, 02000 m. Common.
Note A widespread and polymorphic species.
Macaranga intonsa Whitmore, Kew Bull. 34 (1979) 603; in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 148; Gen. Macaranga Prodr. (2008) 160. Type: Hoogland & Craven 10911 (holo K; iso A, BRI, BO, CANB, G, L, LAE, PNH, US), Papua New Guina, Sepik dist., Sumset (Mt. Hunstein).
Small trees. Indumentum: twigs, stipules and petioles densely pilose with long spreading pale irritant hairs. Twigs 8 mm diam., terete. Stipules ovate, to 6 by 2.5 cm, apex acute, warm brown (in vivo purple), papery, erect, persistent. Leaves: petioles 16 cm, terete, 3 mm diam; blades big, ovate 2536 by 1934 cm, leathery, base broadly shallowly cordate or 5 cm peltate, margin sinuate and minutely distantly denticulate, apex abruptly caudate, 36 cm, drying chocolate brown or dark khaki brown, all nerves including reticulations paler below, very finely and conspicously doubly bullate (viz between the finest nerves), major nerves above and most nerves below rusty velvety with crisped hairs, black granular glandular below, with 3 main and several lesser nerves at petiole insertion, tertiaries and quaternaries scalariform. Staminate inflorescences rusty velvety panicles, 17 by 8 cm, with 4 axis orders; main axis flattened 3 mm diam.; peduncle 5 cm; main branches ascending to spreading, lowest pair opposite, ultimate branches thread-like, zigzag between 3 mm spaced flower clusters; bracts elliptic, variable, to 1cm long, apex acuminate, papery, very soon caducous; bracteoles with a 1 mm broad stalk and spreading ovate, 2 mm diam., obtuse blade, entire, abaxially densely granular glandular and with 5 sunken adaxial glands, fleshy, with a central nerve, spreading, persistent, longer than cluster. Staminate flowers c.10 per cluster, shortly pedicellate; stamens 4, anthers 4-locular. Infructescences 12 cm narrow racemes; peduncle 10 cm, flattened. Fruits 1 cm diam., softly spiny with 2 soon caducous apical sinuous slender lanate 12 mm styles with rigid stigmatic processes. Seeds unknown.
Distribution Northeast New Guinea.
Habitat & Ecology Montane primary and secondary forests, 12002100 m.
Note This species has a general resemblance to M. carrii with which it has in common minutely bullate leaves, in M. intonsa shallowly so and giving the leaves a most attractive appearance. In all details it is amply distinct, though perhaps related.
Macaranga involucrata (Roxb.) Baill., Ιtude Euphorb. (1858) 432; Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1011; Benth., Fl. Austral. 6 (1873) 146; Warburg, Bot. Jahrb. 13 (1891) 352; F.M.Bailey, Queensland Fl. 5 (1902) 145; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 374; L.M.Perry, J. Arnold Arb. 34 (1953) 222; Whitmore, Gard. Bull. Singapore 31 (1978) 56; in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 148; Airy Shaw Kew Bull. 37 (1982) 27; Whitmore, Gen. Macaranga Prodr. (2008) 160. Urtica involucrata Roxb. [Hort. Beng. (1814) 67, nomen nudum] Fl. Ind. ed. 2, 3 (1832) 592. Type: Wallich 4621 (holo K-WALL; iso CGE, G-DC), Malay Islands.
Halecus terrestris Rumph., Herb. Amboin. 3 (1743) 197, 198, t.127, nom. inval.; see Merr., Interp. Herb. Amboin. (1917) 321.
Mecostylis acalyphoides Kurz ex Teijsm. & Binn., Natuurk. Tijdschr. Ned. Indiλ 27 (1864) 44; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 374. Macaranga involucrata (Roxb.) Baill. var. acalyphoides (Kurz ex Teijsm. & Binnd.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1012.
Macaranga mallotoides F.Muell., Fragm. 4 (1864) 139; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 376. Macaranga involucrata (Roxb.) Baill. var. mallotoides (F.Muell.) L.M.Perry, J. Arnold Arb. 34 (1953) 223; Airy Shaw, Kew Bull. 31 (1976) 396; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 149.
Macaranga asterolasia F.Muell., Fragm. 4 (1864) 140, nomen subnudum.
Mappa amboinensis Mόll.Arg., Linnaea 34 (1865) 197. Macaranga amboinensis (Mόll.Arg.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1002.
Macaranga schleinitziana K.Schum., Bot. Jahrb. 9 (1887) 207; K.Schum. & Hollrung, Fl. Kaiser Wilhelms Land (1889) 79; Warb., Bot. Jahrb. 13 (1891) 351; K.Schum. & Lauterb., Fl. Schutzgeb. Sόdsee (1900) 397; Nachtr. (1905) 297; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 373, t. 62FG; L.M.Perry, J. Arnold Arb. 34 (1953) 224.
Macaranga involucrata (Roxb.) Baill. var. keyensis Warb., Bot. Jahrb. 13 (1891) 352. Macaranga keyensis (Warb.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 376.
Macaranga modesta Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 375.
Macaranga schleinitziana K.Schum. var. lobulata Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 374.
Macaranga dalechampioides S.Moore, J. Bot. 61, Suppl. (1923) 48; L.M.Perry, J. Arnold Arb. 34 (1953) 221.
Macaranga seta-felis Whitmore in Airy Shaw, Kew Bull. 36 (1981) 423; Kew Bull. 39 (1984) 608.
Shrubs or small trees, 210 m. Twigs slender, 3 mm diam., terete, finely ochreous pubescent, or occasionally (sub)glabrous. Stipules setose or narrowly elliptic, 812 by 1(2) mm, ochreous tomentose, erect, caducous. Leaves: petioles 38 cm, slender, terete, usually densely (rarely sparsely) ochreous pubescent, some hairs tufted; blades broadly ovate, 1016 by 812 cm, papery to thinly leathery, if the latter occasionally weakly bullate, base usually broadly deltoid, occasionally broadly rounded, 3 mm peltate or closely weakly cordate, with a few inconspicuous glands near petiole insertion margin entire, apex (rarely absent or) caudate, up to 3 cm, very rarely with unequal lateral cusps, drying khaki-brown, above usually with white hairs (a minority tufted) on main nerves near base, below with similar hairs (usually dense) on main nerves, and with dark granular glands, rarely only sparsely hairy; palmately nerved, with 3 main and 2 minor nerves from petiole insertion. Staminate inflorescences 510 cm, dense to open racemes, with distant divaricate often slender branches, distally sometimes thread-like; axes terete, densely finely pubescent; basal peduncle short, 24 cm, rarely with axillary branches; bracts tiny, ovate, 1 mm, apex acute, persistent; flower clusters dense or well spaced, the axis zigzag between them, small, 818-flowered; bracteoles persistent, very variable, tiny, oblong (to ovate), to 2(3) mm long, apex blunt (to acute), usually papery, densely hairy outside or rarely subglabrous, inside with scattered hairs and 15 small inconspicuous patellar glands usually marginal and partly protruding so margin irregular, rarely absent, usually spreading, sometimes recurved, occasionally erect, smaller than cluster. Staminate flowers tiny, globular, 0.5 mm, developing sequentially; pedicel short, thread-like; sepals free, pubescent; stamens 310, anthers 4-locular. Pistillate inflorescences pendulous 510 cm spikes (once a raceme), topped by a cluster of leafy bracts, ovate acute to 15 by 10mm, enclosing c. 5 flowers, sometimes also a group of subapical bracts, axis slender terete, pubescent. Pistillate flowers with calyx tubular, lobed, circumscissile, soon caducous; ovary golden tawny 1(3)-locular with 1(3) subapical, 1015 mm, brittle, plumose, black, free style(s) protruding from bracts. Fruits 1(3)-locular, 3 mm across, closely granular glandular, at first with brittle short spines, wall woody; pedicel 3 mm; calyx persistent; styles brittle, caducous. Seeds round, coarsely verrucose.
Distribution Sulawasi, Moluccas (incl. Kai and Aru Islands), New Guinea (west, southeast), Bismarck Archipelago (New Britain), Austraila (Northern Territory, north Queensland), Solomon Islands, Vanuatu.
Habitat & Ecology Secondary forest, low elevation to 600(1200) m. Very common.
Notes 1. This is a common, widespread polymorphic species. I do not now believe any discrete distinctive varieties can be delineated, even in Australia. It is often a treelet. It is distinctive in its setose to narrowly elliptic stipules, (usually) velvety deltoid leaves with spreading white hairs, at least some of them tufted, and pendulous pistillate spikes (bearing a terminal cluster of leafy bracts). (I have seen only one collection with the pistallate an open raceme, Mueller 56 from Australia, Northern Territory). The staminate inflorescences are variable with tiny flower clusters dense to diffuse, which perhaps changes with age. The staminate flowers are minute and vary in size up to 0.5 mm. The staminate bracteoles are usually densely hairy outside but extremely variable in shape, with the patellar glands small, inconspicuous and rarely absent. An extreme form has ovate, acute, glandless bracteoles but is otherwise indistinguishable. Moreover, there is variation in the bracteoles on a single plant. I am unable to recognise more than one taxon, variable in the staminate parts.
2. When I described M. seta-felis I thought its locality Nusa Laut was an island off Sumatra and because it differs from all other species of the Sunda Shelf I considered it a new taxon. In fact, Nusa Laut is an island south of Seram (Moluccas), and M. seta-felis is none other than M. involucrata, which is widespread and common from Sulawesi eastwards.
3. Provisional reduction of Macaranga modesta and Mappa amboinensis (Ambon: Lesson s.n., which is missing from G-DC) is made on the basis of their descriptions. Macaranga modesta was described from an un-numbered Teysmann collection from Java, which I have to assume was mis-labelled as there is nothing like M. involucrata native to that island.
4. Macaranga asterolasia is sketchily described with no type cited. The weakly peltate leaves with tufted hairs below point strongly to M. involucrata. It is cited and reduced to M. involucrata in Index Kewenis.
5. Macaranga carolinensis replaces M. involucrata in Micronesia.
Macaranga javanica (Blume) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1004; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 340; Whitmore in Airy Shaw, Kew Bull. 36 (1981) 320; Backer & Bakh.f., Fl. Java 1 (1963) 488; . Mappa javanica Blume, Bijdr. (1825) 625; Baill., Ιtude Euphorb. (1858) 430; Zoll., Linnaea 28 (1856) 308, Reichb.f.& Zoll., Verhand. Natuurk. Vereen. Ned.-Indiλ 1 (1856) 9; Whitmore, Gen. Macaranga Prodr. (2008) 162. Rottlera javanica (Blume) Hassk., Flora 25, Beibl. 2 (1842) 41. Tanarius javanicus (Blume) Kuntze, Rev. Gen. 2 (1891) 619. Syntypes: Blume s.n. (L, P), Java, Gunung Gede and Gunung Salak; Kollmann s.n.; Zollinger 652 (G-DC, K, P, W), Java [Welzen: probably only the Blume specimens are the real syntypes, the others are additional material seen by Mόller (1866).]
Mappa bancana Miq., Fl. Ind. Bat., Suppl. (1861) 458. Macaranga javanica (Blume) Mόll.Arg. var.bancana (Miq.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1005.
Small trees. Twigs 3 mm diam., angled at tip, terete further back, irregularly ridged, rusty furfurescent, blackish. Stipules broadly elliptic, 8 mm long, 3 mm wide at base to an acuminate to caudate apex, densely rusty furfuraceous, papery, erect, caducous. Leaves: petioles 57 cm, slender, 1.5 mm diam., terete, weakly furfuraceous; blades ovate oblong, 913 by 47 cm, leathery, base broadly cuneate, sometimes 1 mm peltate, recurved upwards, with 2 petiole insertion glands, margin very weakly sinuate and very weakly recurved, apex acuminate, 1.5 cm, drying dull brown, below sparsely pubescent at least on midrib to nearly glabrous and closely granular glandular. Staminate inflorescences behind leaves, crowded, delicate narrow racemes to 10 cm, rusty pubescent, branches ascending to divaricate; peduncle 2 cm; main axis slender, terete, ridged; lower branches alternate, 2 cm long, usually with axillary branches; bracts and bracteoles somewhat leaf-like, hastate, to 8 by 1.5 mm, narrowed at base, acuminate, margin entire or distantly toothed, with a central nerve, furfuraceous, spreading, persistent; branches thread-like; flower clusters regularly spaced at 8 mm. Staminate flowers in clusters of c. 16, shorter than bracteole, 0.3 mm diam.; pedicel 0.3 mm; sepals 2, granular glandular; stamens 3, anthers 4-locular. Infructescences spikes or narrow sparsely branched racemes, to 14 by 5 cm, as staminate except no axillary branches seen; bracts caducous; bracteoles mostly caducous. Fruits in groups of 13, weakly bilobed, 3 by 4 mm, black granular glandular and with sparse tiny blunt caducous spines, thinly woody; pedicel 34 mm, slender; sepals persistent, recurved; styles 2, apical, small, caducous, recurved. Seeds round, verrucose, with sarcotesta.
Distribution Sumatra (including Bangka), Java.
Habitat & Ecology Lowland to 1300 m.
Notes 1. Sterile material from Java I place here, although leaves cannot be distinguished from M. heynei.
2. It is odd that the staminate and pistillate inflorescences are in different positions.
Macaranga johannium Whitmore, Kew Bull. 34 (1979) 603; in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 149; Gen. Macaranga Prodr. (2008) 163. Type: Henty et al. NGF 33025 (holo K; iso L), Papua New Guinea, Western Distr., near Ingambit.
Small trees, 3 m. Twigs stout, 9 mm diam., slightly angled, with conspicuous stipule scars, glabrous. Stipules ovate oblong, to 8 by 3 cm, apex acuminate, sides slightly curved, warm brown, papery, glabrous, with many close parallel nerves, erect to spreading, persistent. Leaves: petioles 15 cm, strongly channeled, 4 mm diam., weakly scurfy; blades ovate, to 23 by 17 cm, thinly leathery, base broadly shallowly cordate and cordulate at petiole insertion, margin repand with small protruding glandular teeth, apex acute, drying with darker main nerves, lamina fawn above, pale brown below, below with fairly sparse crisped hairs on main nerves and densely dark brown granular glandular, above glabrous, shiny, secondary nerves straight parallel c. 18 pairs, basal pair from petiole insertion, tertiaries scalariform. Staminate inflorescences and flowers unknown. Infructescences stout spikes to 12 cm; axis flattened, 3 mm across, tomentose with long golden hairs; peduncle 5 cm. Fruits crowded towards apex, solitary, weakly 4-lobed, 6 by 14 mm, woody, densely covered with black granular glands and fairly dense slender tapering 5 mm hirsute spines; pedicel 8 mm, stout, sparsely hairy; calyx small, persistent; styles not seen. Seeds unknown.
Distribution East New Guinea (known from type only).
Habitat & Ecology Lowland secondary forest, 140 m.
Note The big papery stipules suggest the Mappa group, of which, however, most species are peltate. I can no longer maintain my Intonsa group.
Macaranga kinabaluensis Airy Shaw, Kew Bull. 23 (1969) 91; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 35; Gen. Macaranga Prodr. (2008) 165. Type: Chew & Corner 4957 (K holo; iso L, SING), Borneo, Sabah, Mt. Kinabalu.
Trees 527 m tall. Twigs to 5 mm diam., angled, glabrous, mid-brown. Stipules narrowly ovate, 5 by 2 mm, rusty pubescent, erect, soon caducous. Leaves: petioles 59 cm, slender, terete, very soon glabrous, not kneed; blades ovate or trullate-ovate, 715 by 49(14) cm, rigidly leathery, base broadly cuneate broadly rounded or broadly truncate, sometimes minutely to 5 mm peltate or cordulate, often with 2 big glands near petiole insertion, margin strongly recurved, apex acute to acuminate, drying pale greenish khaki, maturing glabrous except sparsely hairy on main nerves below, secondaries prominent below, basal secondaries joining midrib at petiole insertion. Staminate inflorescences open panicles to 24 cm long, 3 axis orders, weakly sparsely pubescent; peduncle 4 cm, (slightly) flattened, slender, 1.5 mm diam.; main branches to 7 cm, distant, alternate, without accessories, tertiary branches arising from flower clusters on secondaries; bracts oblong c. 5 by 1 mm, base often subspathaceous, floccose, granular glandular with 14 adaxial patellar glands; some persisting, flower clusters 515 mm apart, subtended by bracteoles; latter linear to subspatulate, 5(10) by 1 mm, furfurescent, with 14 patellar glands, spreading, persistent, with enlarged spathaceous base enclosing the c. 4 flowers. Staminate flowers 1 mm diam. at anthesis, later enlarging, very shortly pedicellate; sepals granular glandular; stamens 4, filament ultimately 1 mm, anthers 4-locular. Pistillate inflorescences amongst and behind leaves, lax slender racemes to 19 cm long, axes slender, sometimes sinuous; peduncle to 6 cm; bracts soon caducous, to 4 by 1 mm, apex blunt, with adaxial patellar glands;bracteoles variable, linear, 13 mm, with 1 or a few patellar glands and finely sparsely furfuraceous, mostly caducous. Pistillate flowers c. 1 cm, apart in clusters of 13; pedicel 13 mm; calyx with 23 unequal lobes. Young Fruits weakly bilobed, depressed, compressed, globose, 3 by 4 mm, closely golden granular glandular and with a few short brittle spines; pedicel 3 mm; calyx minute, persistent; styles 2, apical, recurved, 2 mm, with papillose stigmatic surfaces. Mature Seeds not seen.
Distribution Borneo (Sabah).
Habitat & Ecology Primary forest often near streams, 2001700 m, sometimes on ultrabasics.
Macaranga kingii Hook.f., Fl. Brit. India 5 (1887) 451; Pax & K.Hoffm., Engl., Pflanzenr. IV.147.vii (1914) 383; Ridl., Fl. Malay Penins. 3 (1924) 300; Corner, Ways. Trees Malaya (1940) 302; Whitmore, Mal. Nature Journ. 20 (1967) 98; S.J.Davies, Harvard Pap. Bot. 6 (2001) 411. Type: King's Collector s.n. (holotype: K), Malaysia Johore, Jaffaria.
Macaranga insignis Merr., Philipp. J. Sci. C. Bot. 11 (1916): 69. Lectotype (designated by S.J.Davies, 2001): Native Collector 169 (lectotype: A; isolectotype: K), Malaysia, Sarawak.
Small slender tree to 10 m tall; twigs 713 mm diameter, glabrous, moderately glaucous, hollow, housing ants. Stipules narrowly ovate, 1526 by 813 mm, with a finely acuminate apex, recurved with the margins revolute appearing horn-like and appressed to the stem, glabrous, the pair not completely encircling the twig, succulent, producing food-bodies on abaxial surface, up to 6 pairs present on the shoots. Leaves: petioles terete, 1540 cm long, usually slightly glaucous, glabrous; blades ovate to broadly ovate, 2055 by 2040 cm, adaxial surface dark shiny green, appearing slightly bullate, very deeply dissected with 57 widely spreading lobes, deeply peltate to 410 cm, base very broadly rounded or truncate and sometimes slightly undulate, margin entire, with small protruding flattened nectaries, apices finely acuminate, adaxial surface glabrous, abaxial surface glabrous or sometimes with minute silvery crisped hairs on the main veins and sometimes remaining purple-red when mature, not glaucous; 1° venation palmate with 78 prominent veins, 2° venation looping near the leaf margins and forming a prominent marginal collecting vein, 3° and 4° venation veins weakly scalariform; young leaves dark red below, glabrous or scattered with minute ferrugineus hairs and small yellow glands. Staminate inflorescences paniculate, erect, to 20 by c. 6 cm, glabrous, sometimes slightly glaucous, to 3 axis orders, main axis flattened, first pair of secondary branches ± opposite with or without accessory branches; bracts ovate, 1020 by 812 mm, persistent, margin entire, apex acute, glabrous; flower clusters with 15 flowers, spirally arranged and crowded towards the ends of inflorescence branches; bracteoles ovate, 46 by 34 mm, thin and delicate, enclosing flower clusters, margin entire, apex acuminate, glabrous, scattered with yellow granular glands. Staminate flowers c. 1.5 mm long, pedicellate; sepals free, with scattered minute red-brown hairs and yellow glands, membranous; stamens 47; anthers 4-locular. Pistillate inflorescences paniculate, erect, to 10 cm long, stout, glabrous, usually glaucous, 12 axis orders, secondary branches ± opposite; bracts caducous, not seen. Pistillate flowers c. 12 mm long, solitary in bract axils; calyx urceolate, 46 mm long, not glaucous, with scattered ferrugineus hairs near the base, persistent, splitting irregularly as the ovary expands; ovary 45(6)-carpellate, 46 mm long; styles 49 mm long; styles fused at base, free and spreading from c. 1/2 of the length, persisting to form a prominent 68 mm long crown at the fruit apex; stigma shallowly dissected. Fruits subglobose, 912 by 1015 mm, pale pink to bright red, pedicellate, two discrete glandular patches on each carpel wall developing into narrow ridges raised to 12 mm, covered in dark, sticky exudate; pedicel 1030 mm long, often finely pubescent, not glaucous. Seeds 5.56.5 mm in diam., subtriangular-ovoid, black, with shallow narrow grooves and a small cruciform scar, encased in a fleshy bright orange-red aril.
Distribution Malay Peninsula, Sumatra, Borneo (Sarawak). It has been collected from only three locations: Johore in Peninsular Malaysia, the Tiga Puluh mountains in Sumatra, and between Kuching and Matang in Sarawak. There have been no recent collections of M. kingii from Sarawak, and Davies (2001) has searched extensively in the Kuching and Matang areas and he has not been able to relocate it. Much of the area around Kuching and between Kuching and Matang is no longer covered in intact forest, and the species may no longer exist there. Further exploration in areas close to Kuching is required.
Habitat & Ecology Although Corner (1940) described this species as "one of the more grotesque Malayan plants" it is an extremely elegant small tree often occurring in shaded microsites along streams or in periodically swampy areas.
Note This species is easily distinguished from all other species of Macaranga by the very deeply 57 lobed leaves and strongly recurved horn-like stipules. Macaranga kingii is most closely related to M. umbrosa but differs in having dissected leaves and glaucous stems. It also shares stipule and fruit morphology with M. lamellata, but differs from that species in having glabrous deeply dissected leaves.
Macaranga kostermansii L.M.Perry, J. Arnold Arb. 34 (1953) 251; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 149; Gen. Macaranga Prodr. (2008) 166. Type: Kostermans 2099 (holo A; iso L), Indonesia, Papua, Vogelkop, Anggi Gita Lake.
Shrubs or treelets, to 6 m. Twigs 3 mm diam., terete, ridged, blackish with sparse white hairs. Stipules elliptic, 3 by 1 mm, apex acute, chestnut brown, papery, sparsely hairy without, erect, soon caducous. Leaves: petioles 5 mm, slender, <1 mm diam., terete, ridged, tomentose; blades tiny and crowded, elliptic, 2.54.5 by 11.2 cm, leathery, base cuneate, with a few tiny scattered glands, margin repand, recurved, apex narrowing abruptly to a blunt acuminate caudex, drying khaki, midrib below with sparse white tufted hairs, and densely dark granular-glandular, above midrib raised, keeled and surface weakly shagreened, secondary nerves c. 5 distant pairs, looping near margin. Staminate inflorescences tiny delicate spikes or sparsely branched racemes to 3 cm long; peduncle 03 mm; axes thread-like, <1 mm diam., terete, ridged, granular-glandular and sparsely hairy; branches to 3 mm, spreading, arising from a flower cluster; flower clusters touching or to 3 mm apart; bracteoles ovate, 1 by 1 mm, apex acute, membranous, glabrous, slightly cucullate, margin minutely toothed, spreading, smaller than cluster, persistent. Staminate flowers 6 per cluster, developing sequentially, to 1 mm diam., very shortly pedicellate; stamens 6, anthers 4-locular. Pistillate inflorescences, flowers, fruits, and seeds unknown.
Distribution West New Guinea (Vogelkop, Netoti mountain and Anggi Gita lake).
Habitat & Ecology Forest edges and primary forest, 16002300 m.
Note The type and two other collections are distinctive in their tiny crowded leaves, smaller than in any other species. A fourth collection Versteegh BW 10416, also staminae and also from Netoti, differs mainly in larger (6.5 by 2 cm) papery, non shagreened leaves, and larger, 5 by 2 mm stipules. Clearly more material is needed to complete the circumscription.
Macaranga laciniata Whitmore & Airy Shaw, Kew Bull. 25 (1971) 241; Airy Shaw, Kew Bull. 26 (1972) 290; Whitmore, Biol. J. Linn. Soc. 1 (1969) 226, t. 2; Tree Fl. Malaya 2 (1973) 112; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 373; Whitmore, Gen. Macaranga Prodr. (2008) 168. Type: Whitmore KEP FRI 3813 (holo K; iso SING), Peninsular Malaysia, Pahang, near Tk. Chempadak, Kuantan.
Treelets or trees to 8 m. Twigs and Leaves just like M. heynei. Stipules triangular, c. 10 by 4 mm, acuminate, papery, with fine brown pubescence, recurved, caducous. Inflorescences as M. heynei but shorter and denser, to 6 by 3 cm, always racemose; staminate with alternate branches and axillary branches; staminate flower clusters crowded; ultimate branches arising from a flower cluster; bracteoles ovate, to 8 mm long, deeply finely laciniate sometimes with no central lamina, laciniae with a small apical gland, fairly thick, finely furfuraceous. Staminate flowers clusters of c. 20, 0.2 mm diam.; pedicel, very short; sepals 2, weakly pubescent; stamens 2(3), anthers 4-locular. Pistillate flowers mostly in 3s. Fruits 2(3)-lobed, 3 by 4 mm, slightly flattened, smooth with close dark granular glands; pedicel 23 mm; calyx small, persistent; stigmas tiny, persistent, apical. Seeds smooth, with sarcotesta.
Distribution South Peninsular Thailand, Peninsular Malaysia (E. Pahang, Trengganu, Kelantan).
Habitat & Ecology Lowlands to 360 m.
Note Replaces M. heynei in northeast Malaya, apparently allopatrically, to which it is very similar differing conspicuously in the stipules and bracteoles.
Macaranga lamellata Whitmore, Kew Bull. 29 (1974) 448; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 148; S.J.Davies, Harvard Pap. Bot. 6 (2001) 412. Type: Leopold SAN 74346 (holotype: K), Malaysia Sabah, west of Tampias, Ranau.
Small tree to 15 m tall, flowering from c. 3 m; twigs 815 mm in diam., glabrous, moderately to densely glaucous, hollow, housing ants. Stipules narrowly ovate, 1527 by 813 mm, with a finely acuminate apex, recurved with the margins revolute appearing horn-like and appressed to the stem, glabrous, the pair not completely encircling the twig, succulent, producing food-bodies on abaxial surface, up to 4 pairs present on shoots. Leaves: petioles terete, stout, 1550 cm long, up to 810 mm in diam., usually glaucous, glabrous; blades ovate, often huge, 2065 by 1436 cm, entire or rarely with very short lateral cusps, 410 cm peltate, base broadly rounded, margin entire, with small protruding flattened and conical nectaries, apices acute, adaxial surface mostly glabrous, abaxial surface sparsely to densely scattered with long silvery wavy hairs, moderately to densely glaucous; 1° venation palmate with 79 prominent veins, 2° venation looping near the leaf margins and forming a subprominent marginal collecting vein and also ending at the margin in small flattened protruding nectaries, 3°4° venation scalariform, 5° venation loosely reticulate; young leaves reddish-brown with loose wavy hairs on abaxial surface. Staminate inflorescences paniculate, erect, 412 by c. 6 cm, glabrous, up to 3(4) axis orders, main axis flattened, first pair of secondary branches ± opposite with accessory branches; bracts ovate, 815 by 510 mm, persistent, margin entire, apex acute to acuminate, glabrous; flower clusters with c. 5 flowers, spirally arranged and crowded towards the ends of the inflorescence branches; bracteoles ovate, 46 by 34 mm, enclosing flower clusters, margin ± entire or rarely with 12 cusps, apex acuminate, glabrous or sometimes with very few scattered minute red-brown hairs and yellow granular glands. Staminate flowers c. 1.5 mm long, pedicellate; sepals free, glabrous or with scattered yellow glands, membranous; stamens 45; anthers 4-locular. Pistillate inflorescences paniculate, erect, 514 by 36 cm, stout, glabrous, sometimes glaucous, up to 2 axis orders, secondary branches ± opposite; bracts caducous, not seen. Pistillate flowers solitary in bract axils; calyx urceolate, sometimes glaucous, glabrous, persistent, splitting irregularly as ovary expands; ovary 45(6)-carpellate; styles fused at base, free and spreading from c. 3/4 of the length, persisting to form a prominent 610 mm long crown at fruit apex; stigma not dissected. Fruits subglobose, 912 by 1018 mm, yellowish-green to pale pink to red, sometimes glaucous, pedicellate, two discrete glandular patches on each carpel wall developing into narrow ridges raised to c. 24 mm, covered in yellow, sticky exudate; pedicel 518 mm long, glabrous, usually glaucous. Seeds subtriangular-ovoid, 5.56.5 mm in diam., black, with shallow narrow grooves and a small cruciform scar, encased in a fleshy bright orange-red aril.
Distribution Borneo.
Habitat & Ecology Macaranga lamellata is a relatively shade tolerant tree and is often found within intact primary lowland mixed dipterocarp forest. This species occurs in most areas of Borneo in mixed dipterocarp forest below about 600 m, although it is not nearly as gregarious as the more gap-demanding pioneer species. Macaranga lamellata is commonly found on moderate to low nutrient status sandstone-derived soils. It also appears to be quite tolerant of drought stress. The closely related M. umbrosa has a very similar ecology to M. lamellata and occurs within the same forest formations but that species is more common in forests over more nutrient-rich shale- or basalt-derived soils.
Notes 1. Macaranga lamellata is a member of the small clade of species that includes M. kingii and M. umbrosa. These three species share very distinctive recurved horn-like stipules, staminate flowers with c. 5 stamens, and large fruits on long pedicels with two very prominent narrow glandular ridges per carpel. No other species of Macaranga bear this set of traits.
2. Macaranga kingii is distinguished from M. lamellata in having large deeply dissected five lobed glabrous leaves. Macaranga lamellata differs from M. umbrosa in having glaucous leaves and young shoots, very prominent punctate yellow glands on the lower leaf surface, pubescent mature leaves, the leaf bud with dense pilose hairs, as well as in several features of the flowers and fruits.
3. There is some variation among the specimens Davies (2001) has seen of M. lamellata. The specimens from Lambir Hills National Park in northern Sarawak differ slightly from other areas in being particularly densely glaucous and the abaxial leaf surfaces are somewhat less densely pubescent than collections from other areas.
Macaranga lanceolata Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv, add. VI (1919) 25; L.M.Perry, J. Arnold Arb. 34 (1953) 251; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 149; Gen. Macaranga Prodr. (2008) 169. Syntypes: Ledermann 9030 (K), 9074 (K), 9270 (K), 9496 (K), 9567 (K), 9912 (K), 9942 (K), 10250 (K), all Papua New Guinea, Sepik region, Etappenberg.
Small trees 10 m. Twigs slender, 2 mm, slightly angled, glabrous, often blackish brown. Stipules tiny, elliptic, 3 mm, apex acute, blackish brown, papery, glabrous, erect, caducous. Leaves: petioles 23 cm, slender, 1 mm diam., terete, glabrous or with a few spreading hairs, kneed at top; blades (narrowly) oblong elliptic 512 by 1.53.5 cm, thinly leathery, base cuneate, abruptly narrowing to distinctive 3 mm basal gland-bearing extension, margin repand slightly recurved, apex acuminate, dark brown, glabrous and without granular glands, basal secondary nerves marginal. Staminate inflorescences mostly amongst leaves, crowded short 4 cm racemes, glabrous or slightly furfuraceous; branches 1.5 cm, ascending, thread-like; peduncle 1.5 cm; main axis slender, 1 mm, terete, ridged; flower clusters contiguous or 2 mm spaced; bracteoles deltoid, 0.7 mm, apex acute, entire, spreading, persistent, smaller than clusters. Staminate flowers in clusters 45, sessile; stamens 4, anthers 4-locular. Pistillate inflorescences as staminate. Pistillate flowers solitary; pedicel 35 mm; calyx and ovary furfuraceous; style 2 mm, papillose. Fruits tiny, globose, 3 mm, occasionally bilobed, thinly woody, furfuraceous (not granular glandular); pedicel to 5 mm; calyx persistent. Seeds globose, coarsely rugulose, with sarcotesta.
Distribution East New Guinea (Sepik).
Habitat & Ecology Forest edges, 8501500 m.
Note Like M. faiketo the leaves and fruits lack granular glands and the latter are furfuraceous. The leaf base is distinctive.
Macaranga leytensis Merr., Philipp. J. Sc. Bot. 7 (1912) 393; Enum. Philipp. Flow. Pl. (1923) 442; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 393; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 34; Whitmore, Gen. Macaranga Prodr. (2008) 169. Type: Ramos 15253 (iso K), Philippines, Leyte, Mt. Ibumi, near Dagami.
Trees 12 m. Twigs not seen. Stipules not seen. Leaves: petioles stout, c. 30 cm, round, smooth, 5 mm diam., sparsely rusty tomentose; blades ovate, 45 by 30 cm, papery, base truncate to broadly rounded, narrowly cordate at petiole insertion, margin with distant protruding glands, apex acuminate to 1.5 cm caudate, drying brown, darker above, surface eglandular, and sparse white hairs on nerves below. Staminate inflorescences open panicles to 25 cm long, 4 axis orders, divaricate; main axis rounded, striate, slender, 2 mm diam., tapering, subglabrous, higher orders slightly flattened, rusty furfuraceous; 4th order thread-like with crowded flower clusters; bracts persistent, tiny, ovate, 1 mm; bracteoles persistent, ovate, small to 2 mm across, deeply bluntly regularly toothed, finely furfuraceous. Staminate flowers c. 4 per cluster, tiny, 0.3 mm, pedicellate; sepals free; stamens 2, anthers 4-locular. Pistillate unknown.
Distribution Philippines (known from the type only).
Macaranga lineata Airy Shaw, Kew Bull. 23 (1969) 104; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 150; Gen. Macaranga Prodr. (2008) 170. Type: Hoogland 9199 (holo K; iso A, BM, BO, BRI, CANB, K, L, LAE, NY, US), Papua New Guinea, Morobe Dist., Huon Peninsula, E slope of Mt Rawlingon, along Gang Creek.
Small trees, 6 m. Twigs 7(15) mm diam., coarsely angular, glabrous. Stipules ovate-oblong, 25 by 7 mm, base auriculate, regularly tapering to acuminate tip, with a central nerve, midbrown, papery, glabrous, erect to spreading, subpersistent. Leaves: petioles to 45 cm, to 5 mm diam., terete, coarsely ridged, glabrous, drying constricted at base; blades big, ovate, to 44 by 36 cm, thinly leathery, base broadly rounded, deeply 11 cm peltate, margins entire, apex caudate, 5 cm, drying greenish khaki above, midbrown below, soon glabrous above except for a few hairs on main nerves, below very shortly pilose on nerves and with no granular glands, spider's web nervation, tertiaries and quaternaries raised and conspicuous below invisible above scalariform. Staminate inflorescences 18 by 5 cm narrow racemes, main axis flattened, ridged, 2 mm diam., weakly pilosulose or glabrous; peduncle 4 cm; branches more or less ascending to 4 cm long, puberulous, arising from a flower cluster, basal pair opposite, them and some others with axillary branches; a few persistent variable stipule-like bracts to 1 cm long; bracteoles ovate, to 8 by 8 mm, acute or acuminate, strongly cucullate, papery, conspicuous dark parallel nerves, granular glandular both sides and pubescent, margins closely pectinate the teeth ending in 1 mm hairlike tips, spreading, persistent, loosely enclosing and larger than regularly spaced clusters of flowers. Staminate flowers in clusters of 1520; pedicel 2 mm; stamens c. 8, anthers 4-locular. Pistillate unknown.
Distribution East New Guinea (known from type only).
Habitat & Ecology Secondary forest, 1500 m.
Macaranga loheri Elmer, Leafl. Philipp. Bot. 2 (1908) 432; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 341, t. 55; Merr., Enum. Philipp. Flow. Pl. (1923) 442; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 34; Whitmore, Gen. Macaranga Prodr. (2008) 170. Type: Elmer 8558 (isotypes BO, K, L), Philippines, Benguet Prov., Baguio.
Small trees, 6 m. Twigs slender, terete, glabrous. Stipules elliptic, 8 mm, papery, glabrous, soon caducous. Leaves: petioles 45 cm, slender, terete, glabrous; blades ovate, 714 by 47 cm, papery, base truncate to broadly cuneate, margin rarely crenate, apex acute to acuminate, glabrous, with very fine granular glands below. Staminate inflorescences crowded, to 12 cm with 1 or 2 spreading sparse branches, 2 cm long; axes slender, glabrous; flower clusters 3 mm, in groups; bracteoles conspicuous, persistent, spreading, ovate, to 10 by 1.5 mm, tapering to base, apex caudate, with 2 or 3 big adaxial patellar glands, glabrous. Staminate flowers c. 20 per cluster, sessile, 0.8 mm diam.; calyx divided to base; stamens 3; anthers 4-locular. Pistillate unknown.
Distribution Philippines (Camiguin, Luzon, Samar), Sulawesi (Menado).
Note The conspicuous persistent bracteoles are highly distinctive. When pistillate material is found and if the bracteoles are found to be caducous, then M. loheri can be distinguished from M. cumingii and M. sylvatica by its glabrous (but unfortunately caducous) stipules.
Macaranga longicaudata L.M.Perry, J. Arnold Arb. 34 (1953) 233; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 150; Gen. Macaranga Prodr. (2008) 171. Type: Carr 15303 (holo A, iso K, L, SING), Papua New Guinea, Northern Prov., main range NW of The Gap.
Trees, to 18 m. Twigs slender, 3 mm diam., terete, coarsely rusty pubescent. Stipules elliptic, to 6 by 2 mm, apex acuminate, blackish, papery, at first pubescent, erect, soon caducous. Leaves: petioles c. 3 cm, slender,1.5 mm diam., terete, pubescent, kneed at both ends; blades ovate to deltoid, 710 by 47 cm, leathery, base truncate sometimes very broadly very shallowly cordate, usually with 410 blister-like glands protruding from upper surface at petiole insertion, basal margin strongly recurved upper margin with distant protruding glandular teeth, apex caudate, 1(4) cm, drying blackish brown or rusty below, grey above, strongly finely bullate, each eyelet blistered, above glabrous, below with white straight or crisped single or clustered hairs on all nerves, at first floccose, densely black granular glandular, with 3 main basal nerves. Staminate inflorescences delicate triangular 3-axis panicles, 46 by 45 cm plus basal peduncle of 36 cm, pubescent; secondary axes divaricate to 3 cm; tertiaries to 1 cm, arising from a flower cluster; peduncle terete, 1 mm diam., densely to diffusely coarsely pubescent; bracteoles ovate, 1 by 0.6 mm, apex broadly acute, or deeply coarsely 3-lobed, cucullate, erect, papery, granular glandular outside, smaller than cluster, persistent; flower clusters irregularly crowded. Staminate flowers in clusters of 89, globular, 1 mm, granular glandular; stamens 4, anthers 4-locular. Fruits with slightly eccentric single style.
Distribution East New Guinea.
Habita & Ecology Mountain forests, 24002800 m.
Note Similar to M. induta but easily told apart by the leaf base and strong bullation.
Macaranga longistipulata (Kurz ex Teijsm. & Binn.) Mόll.Arg. in DC., Prodr. 15, 2 (1866)991; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 369; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)134 passim; Airy Shaw, Kew Bull. 37 (1982) 27; Whitmore, Gen. Macaranga Prodr. (2008) 172. Rottlera longestipulata Kurz ex Teijsm. & Binn., Natuurk. Tijdschr. Ned.-Indiλ 27 (1864) 43. Type: Teysmann 5193 (holo BO; iso G), Moluccas, Ternate.
Trees to 20 m. Twigs slender, round, blackish, glabrous, slightly pruinose, with encircling stipule scars. Stipules united, apparently as a single terminal elliptic slightly sinuous bud, 512 cm long, caducous, papery, lanate with long adpressed grey brown hairs. Leaves: petioles 2 cm, slender, round, striate, usually glabrous, or with long golden adpressed hairs; blades elliptic, 1524 by 46 cm, thinly leathery, base cuneate cordulate, margin shallowly sinuate, apex long tapering to acuminate, drying grey green above, mid brown below, with sparse adpressed golden hairs on midrib and secondary nerves below, closely golden granular glandular below, midrib raised above but often set in a narrow trough, secondary nerves conspicuous, close-set, parallel, straight, steeply ascending, 12 >20 pairs. Staminate unknown. Infructescences spikes to 5 cm long with a single terminal fruit; axis slender, flattened, with caducous golden hairs and bracteole scars; bracteoles opposite, caducous, elliptic, 10 by 1 mm, including 5 mm stalk, with caducous golden hairs on stalk and abaxial surface, and c. 5 patellar glands on adaxial surface, margin sinuate. Fruits depressed roundish, to 8 mm across, 6 mm long, weakly bilobed, closely covered with flat slender slightly hooked spines; pedicel 3 mm; calyx large, 5 mm, 2-partitie, caducous; style single, apical, 8 mm, soon caducous, dividing into 2, 1 cm long barbed stigmas. Seeds round, large, 5 mm diam., with sarcotesta, coarsely tuberculate.
Distribution Moluccas (Morotai, Ternate, Seram).
Habitat & Ecology Montane forests, 10001500 m.
Note A big montane tree, highly distinctive in its stipule, leaf nervation and fruit ornamentation.
Macaranga louisiadum Airy Shaw, Kew Bull. 23 (1969) 105; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 150; Gen. Macaranga Prodr. (2008) 173. Type: Brass 28065 (holo K), Papua New Guinea, Milne Bay Prov., Louisiade Archipelago, Sudest Island, Rambuso.
Small trees, to 10 m. Twigs slender, 23 mm diam., slightly angled, finely pubescent. Stipules subulate, 6 mm, erect or somewhat recurved, pubescent, subpersistent. Leaves: petioles 23 cm, slender, 12 mm diam., terete, pubescent, slightly swollen at base; blades ovate-elliptic, (8)13(15) by (4)6(9) cm, papery, base rounded, often minutely shallowly cordulate, often with a few superficial inconspicuous glands near petiole insertion, margin weakly sinuate, apex (usually abruptly) acuminate, drying greyish above, khaki below, glabrous above, below with dark granular glands and pubescent to almost glabrous on lower midrib, basal secondary nerves and several minor nerves arising at petiole insertion. Staminate inflorescences dense panicles, to 13 by 5 cm, with 3 axis orders, (finely) pubescent; branches divaricate; peduncle 12 cm; main axis slender, 1 mm diam., terete, lower branches alternate, a few short axillary branches; bracts ovate, 7 by 6 mm, apex abruptly acuminate, very strongly cucullate, pubescent on nerves, granular glandular, very soon caducous; secondary and tertiary branches thread-like; flower clusters in small (5 mm) tight groups at branch tips; bracteoles ovate, 2 by 3 mm (or more), cucullate, with 15 cuspidate teeth, wholly enclosing cluster, outside sparsely tomentose and with scattered black granular glands, inside glabrous. Staminate flowers 9 per cluster, tiny, 0.2 mm; pedicel, long, slender; sepals free, densely puberulous; stamens 2, anthers 4-locular. Infructescences 9 cm spikes; axis as staminate; bracts not seen. Fruits (immature) mostly in small terminal clusters, strongly bilobed, 5 by 10 mm, thinly woody, sparse black granular glands and caducous short 1 mm stiff finely pubescent spines; pedicel, stout, 1 mm, pubescent; calyx tiny, persistent. Seeds unknown.
Distribution Southeast New Guinea (Milne Bay, D'Entrecasteaux and Louisiade islands).
Habitat & Ecology Secondary lowland rain forest.
Note Variable from softly pubescent to almost glabrous.
Macaranga lowii King ex Hook.f., Fl. Brit. India 5 (1887) 453; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 364; Ridl., Fl. Malay. Penins. 3 (1924) 304; Airy Shaw, Kew Bull. 26 (1972) 290; Alph. Enum. Philipp. Is. (1983) 35; Whitmore, Tree Fl. Malaya 2 (1973) 112; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 154; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 373; Whitmore, Gen. Macaranga Prodr. (2008) 173. Syntypes: Cantley s.n. (not seen), Singapore; Curtis s.n. (not seen), Malaysia, Penang, on Goot Hill; King's Collector s.n. (not seen or 3664, K), Malaysia, Perak; Wallich s.n. (cat. 3664, K-WALL), Malaysia, Penang.
Mallotus auriculatus Merr., Philipp. J. Sc. Bot. 7 (1912) 396; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 194; Merr., Enum. Philipp. Flow. Pl. (1923) 432; Ridl., Fl. Malay. Penins. 5 (1925) 333. Macaranga auriculata (Merr.) Airy Shaw, Kew Bull. 19 (1965) 325; Kew Bull. 26 (1972) 287; Gagnep. in Lecomte, Fl. Indo-Chine 5 1926) 448; Whitmore, Gard. Bull. Singapore 26 1972) 62; Tree Fl. Malaya 2 (1973) 112; Chiu, Guihaia 2 (1982) 150.
Mallotus affinis Merr., Philipp. J. Sc. Bot. 13 (1918) 82; Pax & K.Hoffm. in Engl., Pflanzenr. IV,147,xvii, add. vii (1924) 183.
Macaranga poilanei Gagnep., Bull. Soc. Bot. France 69 (1923) 703; in Lecomte, Fl. Indo-Chine 5 (1926) 448, t 53/711, 54/1; Croizat, J. Arnold Arb. 23 (1942) 51; Airy Shaw, Kew Bull. 21 (1968) 406 passim.
Mallotus tsiangii Merr. & Chun, Sunyatsenia 1(1930) 63.
Macaranga lowii King ex Hook.f. var. kostermansii Airy Shaw, Kew Bull. 23 (1969) 107; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 155.
Macaranga glaberrima sec Airy Shaw, Kew Bull. 19 (1965) 322, pro parte quoad specimina borneλnsia, nec Rottlera glaberrima Hassk.
Small trees, to 20 m, dbh up to 16 cm; bole straight. Bark smooth, shallowly cracked, latex red; inner bark thin, dark red. Twigs solid, round, striate, slender, glabrous. Stipules and bud scales 7 mm long, subulate with a small ovate base, chestnut brown glabrous, stiffly erect, crowded at twig tip, soon caducous. Leaves: petioles 37 cm, slender, 1 mm, terete, weakly pubescent or glabrous, kneed; blades elliptic, 815 by 35 cm, thinly leathery, base rounded to cuneate, minutely auriculate, with 2 glands at petiole insertion, margin entire, slightly sinuate, apex abruptly acuminate, 1.5 cm, drying chestnut brown to grey green, below glabrous or pubescent on lower part of midrib and closely to sparsely but regularly dark granular glandular. Staminate inflorescences often malformed as a witches' broom, slender spikes to 6(11) cm; peduncle 1 cm, axis slender, 1 mm diam., finely pubescent, terete to slightly flattened; bracts not seen; flower clusters 34 mm across, 1 cm spaced; bracteoles persistent, ovate, 1.5 mm, apex acuminate, pubescent both sides, shorter than flower cluster. Staminate flowers 4 per cluster, opening serially; sepals hooded, pubescent outside with granular glands towards tip; stamens 12, anthers 4-locular. Infructescences spikes to 9 cm, axis slender, flattened, pubescent, leafy bracts ovate, to 10 by 5 mm, apex acute, soon caducous. Fruits spaced along top 1/32/3 of spike, solitary with a furthergroup at apex, strongly dicoccous, reaching 6 by 12 by 6 mm, woody, when young very densely spiny, at maturity spines spaced, sometimes very sparse, spines 4 mm, flattened, surface furfuraceous, glabrous or black granular glandula; pedicel stout, 24 mm, pubescent; sepals ovate, 2 mm, apex acute, persistent; styles 15 mm, spreading, soon caducous. Seeds round, 4 mm, shiny, smooth, without sarcotesta.
Distribution China (Hainan, Yunnan), Vietnam (north, central), Thailand (Peninsular and southeast), Malay Peninsula, Borneo, Philippines (Panay, Samar, Leyte, Mindanao).
Habitat & Ecology Slik et al.: Mainly in the understorey of primary forests but also in secondary forests. On well drained, clay-rich and sandy loam soils. Up to 1350 m altitude.
Notes 1. The commonest and most wide ranging of the west Malesian primary forest Macaranga species which form Section Pseudorottlera.
2. I consider this to be a widespread polymorphic species, just as variable as M. andamanica, with which it is partially sympatric and of similar Gestalt (for differences see M. andamanica). The leaves vary from brown to greenish, and glabrous to pubescent. This variation encompasses M. auriculata as I pointed out in 1972 (see above). The fruits are usually spiny but vary in their surface which may be pubescent, glabrous or granular glandular. Airy Shaw's var. kostermansii, distinguished in being glabrous with greenish leaves, cannot be maintained: one syntype of the species, King's Collector 3664 (K), is glabrous!
Macaranga lugubris Whitmore, Kew Bull. 39 (1984) 609; Gen. Macaranga Prodr. (2008) 175. Type: Vinas & Wiakabu LAE 59484 (holo K, iso L), Papua New Guinea, Star Mountains, West Sepik Dist., Telefomin Subdist., Folongonom on track to Busilmin.
Gloomy tree, 12 m tall. Twigs 3 mm diam., terete, striate, glabrous, blackish. Stipules oblong, 10 by 2 mm, apex obtuse, blackish, papery, glabrous, erect, subpersistent. Leaves: petioles 4 cm, slender, terete, glabrous, kneed at both ends; blades broadly elliptic, c. 9 by 4 cm, papery, base broadly cuneate, cordulate with 2 large glands, margin repand, apex acumiante, to 1 cm, dark dull brown, underside densely but minutely granular glandular, at first with a cream scurfy covering. Staminate inflorescences narrow 6 cm sparsely branched panicles with 3 axis orders; lowest branches subopposite, divaricate 2 cm with divaricate 5 mm tertiaries, upper 1 cm unbranched, weakly ochreous scurfy; peduncle 1 cm; main axis slender, 1 mm diam., terete; bracts ovate, 1.5 mm, apex acute, erect, papery, persistent; flower clusters evenly spaced, 5 mm to almost touching; bracteoles persistent, broadly triangular, 2 by 2.5 mm, erect, slightly concave, papery, glabrous, sparsely nerved. Staminate flowers 3 per cluster, big, 1.3 mm diam., shortly stalked; stamens 8, anthers 4-locular. Pistillate unknown.
Distribution East New Guinea (only known from the type).
Habitat & Ecology Secondary montane rain forest, 2300 m.
Note Distinctive in its gloomy appearance (to which the epithet refers), its stipules, leaves very minutely granular glandular and few big flowers per staminate cluster.
Macaranga lumiensis Whitmore, Gen. Macaranga Prodr. (2008) 175. Type: Streimann & Martin LAE 52904 (holo K; iso BRI, CANB, L), Papua New Guinea, West Sepik Dist., Lumi Subdistr., Lumi to Nuku road.
Treelet, 3 m. Twigs 3 mm diam., terete, khaki, tomentose. Stipules ovate, 7 by 3 mm, apex acute or weakly acuminate, papery, tomentose, erect, soon caducous. Leaves: petioles to 4 cm, slender, 1 mm diam., terete, tomentose, weakly kneed at top; blades ovate, c. 13 by 6 cm, papery, base rounded, margin sinuous with sparse, inconspicuous, tiny protruding glands and protruding lamina hairs, apex acute, drying khaki, above midrib densely tomentose with spreading mostly single, a few tufted, straight white hairs and lamina with similar hairs arising from small pustules, below lamina densely golden granular glandular, all nerves including reticulations with white hairs, dense on midrib. Staminate inflorescences behind leaves, crowded narrow delicate sparsely branched racemes to 9 by 3 cm; axes furfuraceous; branches divaricate; peduncle 1 cm; main axis slender, < 1 mm diam., terete, lower branches alternate; bracts not seen; branches thread-like, to 2 cm; compound flower clusters 23 mm apart, evenly spaced, each compound cluster of 23 close bracteoles each subtending 1 flower; bracteoles ovate. 2 by 1.3 mm, apex acute, concave, margin entire to weakly serrate, minutely finely velvety, nerved, erect, persistent. Staminate flowers 1 by 0.3 mm with tubular glabrous calyx; stamens 3, protruding, anthers 4-locular. Pistillate unknown.
Distribution East New Guinea (only from the type).
Habitat & Ecology Secondary forest, 500 m.
Note This fits the Gracilis group, but has unusually large leaves and is a lowland plant. Macaranga misimae, also sometimes in the lowlands, also has hairy leaves.
Macaranga magnifolia L.M.Perry, J. Arnold Arb. 34 (1953) 236; Airy Shaw, Kew Bull. 23 (1969) 95 passim; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 150; Gen. Macaranga Prodr. (2008) 175. Type: Brass 1090 (holo A; iso K, P), Papua New Guinea, Gulf Prov., Purari Delta, Mame River.
Trees to 20 m. Twigs stout, to 1 cm diam., terete, with conspicuous leaf and petiole scars, at first with coarse white hairs, becoming glabrous. Stipules broadly oblong-elliptic, 210 cm long, 0.73 cm wide at base, papery, tapering regularly from base to acuminate tip, drying mid to dark brown, outside with moderately dense coarse white to golden hairs inside glabrous, inconspicuously striate, forming an erect bundle protruding beyond twig tip and also persistent along twig. Leaves: petioles 50 cm, stout, 8 mm diam., flattened above at base, terete upwards, with sparse coarse hairs especially at top to glabrous, kneed at top; blades big, ovate-oblong, 2467 by 1660 cm, papery or occasionally leathery, base broadly rounded, 417 cm peltate, margin repand sometimes with distant tiny protruding glands, apex (abruptly) acuminate, 2 cm, drying khaki to dark brown, below the nerves sometimes darker, above glabrous or puberulous on main nerves, below with sparse coarse hairs at least on main nerves near petiole insertion and with tiny (fairly) sparse black granular glands, spider's web nervation, tertiaries and quaternaries scalariform. Staminate inflorescences big open panicles, to 40 by 16 cm, with 4 axis orders; lower axes glabrous, distally rusty furfuraceous; secondary axes ascending, other axes divaricate; peduncle 26 cm, somewhat flattened, unevenly sparsely ridged, to 4 mm diam., sometimes basally with bract scars; main axis stout, tapering, branches mostly alternate, a few opposite, a few axillary branches, secondaries to 13 cm, tertiaries (to 3 cm) and quaternaries (1 cm) bearing contiguous regularly displayed spirally arranged flower clusters; bracts mostly caducous, leaving scars, ovate, 3 by 2 mm, apex broadly acute, rusty scurfy to almost glabrous; bracteoles persistent, ovate, apex rounded, margin dentate to entire, strongly concave, both sides finely densely pubescent, erect, wholly enclosing flower cluster, at first 2 by 2 mm, thick, becoming at anthesis 5 by 4 mm, thin, nerved, the nerves ending at small glands on upper part of margin. Staminate flowers 11 per cluster, oblong, 0.4 mm long, sessile; calyx fused, pubescent; stamens 3, anthers 4-locular. Infructescences stout dense pubescent racemes to 12 by 7 cm; peduncle 34 cm, flattish, 1 cm diam. at base, bearing bracts; latter close, soon-caducous, ovate, 5 by 5 mm, apex acute, concave, pubescent; main axis tapering, branches to 3 cm, ascending, stout, 1.5 mm diam. Fruits solitary, contiguous as a dense mass, with 2 globose lobes, slightly flattened, 5 by 14 by 4 mm, woody, with dense black granular glands and close to sparse, slightly protruding, tawny scurfy round warts; pedicel 2 mm, stout, tawny scurfy; calyx unlobed or trilobed. tawny scurfy. persistent; stigma plate-like, apical, tiny, 1 mm diam. Seeds globose, 4 mm, irregularly shallowly sculptured, with sarcotesta.
Distribution New Guinea (mainly east), Solomon Islands (New Georgia, Kolombangara, Ranongga).
Habitat & Ecology Lowland rain forest, usually primary, sometimes swamps.
Note See notes at M. fragrans.
Macaranga mappa (L.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1000;. Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 320, t. 54A; Merr., Interp. Herb. Amboin. (1917) 29, 319; Airy Shaw, Kew Bull. 23 (1969) 94 passim; Alph. Enum. Philipp. Is. (1983) 27; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 136 passim; Gen. Macaranga Prodr. (2008) 180. Folium mappae Rumph., Herb. Amboin. 3 (1743) 172, t. 108, nom. inval. Ricinus mappa L. in Stickm., Herb. Amboin. (1754) 4; Amoen. Acad. 4 (1759) 124; Sp. Pl. ed. 2 (1763) 1430. Acalypha mappa (L.) Willd., Sp. Pl. 4 (1805) 526. Neotype (Merrill, 1917): Robinson Pl. Rumph. Amb. 361 (K), Indonesia, Moluccas, Ambon, Wae.
Mappa moluccana Spreng., Syst. Veg. 3 (1826) 878.
Macaranga gigantifolia Merr., Philipp. J. Sc. 7 (1912) 391; Enum. Philipp. Flow. Pl. (1923) 441; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 318; Airy Shaw, Kew Bull. 23 (1969) 94 passim; Alph. Enum. Philipp. Is. (1983) 26.
Usually small trees to 10(25) m. Twigs stout, 15 mm diam., terete, furfurescent to pubescent soon becoming glabrous. Stipules big, oblong-elliptic, 4.5(12) by 2(3) cm, widest at base, apex acute to acuminate, warm brown, occasionally slightly glaucous, papery, pubescent with long silky hairs, becoming glabrous, persistent, erect. Leaves: petioles stout, to 50 cm long, to 1 cm diam., angled but not flattened above, glabrous or pubescent especially at apex; blades huge, broadly ovate, 60 by 40 cm, leathery, base broadly rounded, deeply 12 cm peltate, sometimes with small basal glands, margin entire, apex broadly acute, warm brown, pubescent on main nerves above, below with short tufted hairs on nerves to almost glabrous and with dark fairly close granular glands, sometimes with the minor nerves strongly raised below. Staminate inflorescences big, eventually rather open panicles to 50 cm long, 3 axis orders, ochraceous furfurescent; main axis stout, tapering, 5 mm diam. at base, angular; basal peduncle to 6(28) cm, basal part with close caducous bracts leaving conspicuous scars, other axes flattened, sometimes with axillary branches, secondaries divaricate, basal ones 5 cm apart, tertiaries close, 1 cm or less; bracts on main axis oblong, to 10(30) by 5(10) mm, papery, soon caducous, leaving a conspicuous scar, on other axes subpersistent, tiny, ovate, 2 mm, apex acute; flower clusters very regularly displayed in a close variously tight spiral; bracteoles persistent, erect, ovate, 3 mm long by 4 mm broad, apex rounded, slightly concave, papery, shallowly finely crenate, finely furfuraceous, with scattered granular glands. Staminate flowers 720 per cluster, minute, 0.3 mm diam.; sepals free or fused at base; stamens 2, anthers 4-locular. Pistillate inflorescences narrow, 14 cm, ochraceous furfurescent racemes, with adpressed to slightly spreading 23 cm alternate robust branches; peduncle only 1 cm; main axis terete; basal bracts caducous leaving conspicuous scars, others persistent, small, ovate, 1 mm; flower clusters close, evenly spaced, subtending bracteoles as staminate. Pistillate flowers with sepals joined at base. Infructescences furfuraceous; main axis stout, 4 mm diam., strongly tapering; branches fairly stout, tapering to 3 cm; basal peduncle to 2 cm with close bract scars and a few subpersistent, ovate, 11 by 6 mm bracts; bracteoles usually with a short broad stalk, blade ovate, to 5 by 3 mm, apex acute, margin finely crenate, furfuraceous, mostly caducous. Young fruits solitary, at first with close oblong, fine granular glands and scattered conspicuous warts; stigmas plate-like with irregularly frilled margin; fruits later becoming (1)2(3)-lobed, subsessile; mature fruits close, strongly dicoccous, to 10 mm broad by 5 mm deep, woody, closely granular glandular; pedicel stout, 1 mm; calyx small, persistent, unlobed; styles free, 12 mm, finely feathery, breaking off. Seeds spherical, finely verrucose.
Distribution Philippines (Luzon, Samar, Mindanao), Talaud, throughout Sulawesi (Taliabu, Mangole), Moluccas (Halmahera, Morotai, Bacan, Ambon, Seram, Aru, Kai).
Habitat & Ecology Common.
Notes 1. Similar when sterile to M. grandifolia (see there).
2. The tufted white hairs on the leaves below and the infloresceneces with stout main axes and a bare basal bract-bearing peduncle are distinctive. There is considerable variation in hairiness of stipules and leaves and some plants are almost glabrous. The topotype from Ambon (Robinson 361) has very young male inflorescences. Both M. gigantifolia (which I here reduce) and M. mappa were described just from staminate material. Fruiting numbers have been placed here on leaf and stipule characters. Pistillate flowers remain unknown. Rumphius' description and plate clearly show a tricoccous fruit and the plate shows smaller stipules. Despite these differences I follow Merrill's interpretation as I have seen nothing closer to the Rumphian description.
Macaranga melanosticta Airy Shaw, Kew Bull. 25 (1971) 539; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 150; Gen. Macaranga Prodr. (2008) 183. Type: Brass 30745 (holo K; iso L), Papua New Guinea, Eastern Highlands Distr., Mt. Wilhelm.
Trees, 620 m tall. Twigs 3 mm diam., slightly angled, ridged, patchily furfuraceous, becoming glabrous. Stipules elliptic, to 10 by 3 mm, apex acute, dark brown, papery, slightly furfuraceous, erect, soon caducous. Leaves: petioles 2(4) cm, 2 mm diam., terete, ridged, granular glandular, kneed at both ends; blades ovate-oblong, 712 by 46 cm, rigidly leathery, base broadly rounded, very shallowly and broadly cordulate, with 3 basal and sometimes some minor nerves, margin slightly repand, apex fairly abruptly acuminate, 1 cm, drying chocolate brown below, usually slightly concave and weakly coarsely bullate between secondaries, glabrous, slightly shining above, with dense black granular glands below, secondaries and reticulations raised on both surfaces. Staminate inflorescences sparsely branched 810 cm glabrous racemes; branches divaricate, to 2 cm; long peduncle to 4 cm; main axis stout, flattened, striate; branches alternate, large, 5 mm diam.; flower clusters irregularly 3 mm spaced; bracts elliptic, 7 by 3 mm, apex acute or acuminate, papery, sparsely pubescent, soon caducous; bracteoles ovate, 4 by 4 mm, apex acute, broadly shouldered, strongly concave, papery, faintly nerved, granular glandular without, spreading, persistent, smaller than 8-flowered clusters. Staminate flowers globose, 2 mm in bud, opening to 34 mm, sessile; sepals papery granular glandular; stamens 9, anthers 4-locular. Infructescences spikes to 6 cm; peduncle to 4 cm, terete, granular glandular. Fruits 14 per infructescence, globose, to 6 mm diam., leathery, finely pubescent and black granular glandular; pedicel stout. eventually 1 cm; calyx persistent; styles 2, apical, 35 mm, persistent, smooth. Seeds slightly rugulose, with sarcotesta.
Distribution New Guinea.
Habitat & Ecology Secondary or primary montane forest, 24003300 m.
Macaranga minahassae Whitmore, Gen. Macaranga Prodr. (2008) 186. Type: Koorders 16902ί (holo L; iso BO), Sulawsi, Menado.
Small tree. Twigs stout, 15 mm diam., round, very finely pubescent and with spreading fugaceous white hairs, (sometimes) glaucous. Stipules triangular to oblong triangular, 2050 by 1520 mm, apex acuminate, dark brown, ochreous pubescent, papery, erect, persistent (red-green or cream in vivo). Leaves: petioles c. 35c m, stout, to 5 mm diam., tapering, round, with fugaceous spreading white hairs; blades big, ovate, to 45 by 37 cm, papery, base broadly rounded, 612 cm peltate, apex acute, drying brittle, below granular glands present, sometimes sparse, inconspicuous, nerves finely pubescent with crinkled white hairs. Staminate inflorescences narrow sparsely branched, axillary racemes, to 25 cm; basal peduncle 9(15) cm, round, striate, furfuraceous; branches flattish, secondaries slender to thread-like, sinuous, c. 4 cm, all usually with axillary branches, sometimes later developing short tertiaries; bracteoles ovate, 35 by 3 mm, apex acuminate, persistent, strongly concave, irregularly dentate to crenate, with inconspicuous marginal glands, nerved, furfuraceous, wholly enclosing flower cluster. Staminate flowers in spaced clusters of 67, 0.30.5 mm; pedicel 0.30.5(1) mm; stamens 47; anthers 4-locular. Pistillate inflorescences and flowers unknown. Fruits (1 seen) squarish, 4-locular, 10 by 6 by 6 mm, with sparse 3 mm spreading spines; pedicel 8 mm; calyx tiny, persistent; stigmas minute, apical, persistent. Seeds enclosed in black sarcotesta.
Distribution Sulawesi.
Note This is superficially similar to M. leytensis but with smaller, depauperate staminate inflorescences and spiny fruits. Its alliance lies with M. tanarius, note the similarly nerved bracteoles and the tendency for late development of inflorescence axes from within a flower cluster.
Macaranga misimae Airy Shaw, Kew Bull. 23 (1969) 109; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 151; Gen. Macaranga Prodr. (2008) 186. Type: Brass 27533 (holo K; iso L), Papua New Guinea, Milna Bay Prov., Louisiade Archipelago, Misima Island, Mt. Sisa.
Shrubs or small trees, to 15 m. Twigs slender, 23 mm diam., terete, densely softly tomentose with somewhat crisped often tufted hairs. Stipules elliptic, 4 by 1 mm, apex acuminate, chestnut brown, papery, both sides sparsely to densely tomentose, erect, subpersistent. Leaves: petioles 510 mm, slender, 1 mm diam., terete, densely tomentose; blades elliptic to oblong-elliptic, 89.5 by 23 cm, thinly leathery, tapering gradually to a finely cordate base with 27 small glands in lobes and an acuminate apex, margin sinuate to subcrenate, drying khaki brown, above with sparse white hairs, below main nerves densely tomentose with spreading white hairs some tufted and densely black granular-glandular, secondary nerves 1113 pairs, well-spaced, looping and joining near margin, lower part of lamina often somewhat bullate. Staminate inflorescences small delicate sparsely branched 37 cm racemes with dark granular glands and long dense spreading tufted white hairs; branches alternate, arising from a flower cluster, ascending, thread-like, 3 cm; peduncle 1 cm, terete, slender (< 1mm); flower clusters 3 mm evenly spaced; bracteoles oblong, 3 by 1 mm, apex acute, margin sinuous, fleshy, outside dark granular-glandular, within with 2 submarginal rows each of 4 patellar glands, spreading, persistent, longer than flower cluster. Staminate flowers 68 per cluster, minute, 0.3 mm, sessile; calyx granular-glandular; stamens 3, anthers 3-locular. Infructescences delicate spikes to 3 cm with 1(2) apical fruits; axis slender (< 1mm), densely tomentose; bracteoles as staminate but erect, soon caducous. Fruits globose, 5 mm, leathery, sessile, finely black granular-glandular and at first densely beset with tiny obconical hair-tipped knob; calyx persistent s; stigma lateral, 23 mm, papillose, soon caducous. Seeds ellipsoid, globose, rugulose, with sarcotesta.
Distribution New Guinea (Biak island, Eastern Highlands, Misima island).
Habitat & Ecology Primary and secondary forest, sealevel to 1460 m.
Note Known from only three widely separate localities. This could be a form of M. advena (see there).
Macaranga motleyana (Mόll.Arg.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 994; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 310; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 148. ; Kew Bull. 36 (1981) 317; S.J.Davies, Harvard Pap. Bot. 6 (2001) 413; Whitmore, Gen. Macaranga Prodr. (2008) 18, Fig. 13. Mappa motleyana Mόll.Arg., Flora 47 (19864) 467. Type: Motley 1215 (holotype: K), Indonesia, South Kalimantan, Bangarmassing [sic].
Slender glaucous tree to 20 m tall with branches clustered towards the ends of the axes, flowering from 45 m tall in open areas; twigs 69(10) mm diameter, glabrous, rarely with some scattered minute ferrugineus hairs towards the apex, densely glaucous, hollow, housing ants, usually exuding some reddish clear latex when cut. Bark smooth, hoop marked, mottled pale grey-brown and white. Stipules rather small, ovate, 47 by 67 mm, the pair not completely encircling the twig, recurved, succulent, adaxial surface sometimes with scattered minute ferrugineus hairs when young, abaxial surface producing food-bodies, 24 pairs present on the shoots but often caducous. Leaves: petioles terete, slender, 1020 cm long, glaucous, glabrous or sometimes with scattered minute ferrugineus hairs; blades ovate, (15)2035 by (10)1225 cm, usually deeply trilobed, sometimes with an additional pair of basal leaf cusps or lobes, dissected to 1/31/2 of the leaf length rarely less, central lobe c. 48 cm wide usually quite narrow near the base, often with ± parallel sides, lateral lobes c. 37 cm wide ascending and remaining quite close to the main lobe with narrow sinuses, 25(6) cm peltate, base characteristically narrowly rounded, margin entire, apices narrowing suddenly, shortly acuminate, adaxial surface glabrous, abaxial surface mostly glabrous but usually with minute ferrugineus hairs on the veins, densely glaucous; 1° venation palmate with 67(8) prominent veins, 2° venation scalariform, looping near the leaf margins and terminating at the margin in tiny protruding conical nectaries, 3°4° venation only feintly visible not prominently raised from the abaxial leaf surface; young leaves red-brown, minutely ferrugineus pubescent on the adaxial surface and densely yellow granular glandular on the abaxial surface, soon becoming glabrous. Staminate inflorescences paniculate, erect, 1022(25) by 1018 cm, basal axes mostly glabrous and densely glaucous, distal axes sparsely to densely ferrugineus pubescent, up to 4 axis orders, main unbranched axis 28 cm long flattened, first pair of secondary branches ± opposite with accessory branches; bracts elliptic, 717 by 36 mm, usually persistent, bright yellow-green when fresh and drying black, margin entire, apex acute, minutely ferrugineus pubescent; flower clusters with c. 10 flowers, decussately arranged and strongly grouped on ultimate inflorescence branches; bracteoles ovate to broadly ovate, 35 by 2.54 mm, enclosing flower clusters, margin quite deeply and evenly pectinate with c. 14 equally spaced teeth, apex acute and usually terminated by a narrowly caudate apex c. 1 mm long with a tiny conical nectary, both surfaces scattered to densely covered with minute silvery to golden red-brown hairs and yellow granular glands, adaxial surface with a dense patch of minute ferrugineus hairs at the base. Staminate flowers c. 1 mm long, shortly pedicellate; sepals free, splitting irregularly to the base at anthesis, apex usually scattered with minute red-brown hairs; stamens 23; anthers 4-locular. Pistillate inflorescences paniculate, erect, 410 by 35(6) cm, stout, covered in fine ferrugineus hairs, fewer hairs and more glaucous towards the base, 2(3) axis orders, secondary branches ± opposite without accessory branches; bracts caducous, not seen. Pistillate flowers c. 57 by 45 mm, solitary in bract axils; calyx urceolate, 34 mm long, densely covered with fine ferrugineus hairs and scattered yellow granular glands, persistent, splitting irregularly as ovary expands; ovary 45(6)-carpellate, c. 3 mm long; styles c. 23 mm long, fused at base, free and spreading from c. 1/2 of the length, persisting to form a prominent 35 mm long crown at the fruit apex; stigma not dissected. Fruits subglobose, compressed, 79 by 913 mm, pedicellate, sutures bluish and glaucous, one discrete glandular patch on each carpel wall developing into a slightly raised flat disc covered in yellow, sticky exudate; pedicel 48 mm long, scattered with minute ferrugineus hairs, glaucous. Seeds subtriangular-ovoid, 3.54 mm in diam., black, with shallow coarse grooves and a small cruciform scar, encased in a fleshy bright red aril.
Distribution Borneo (Central, E. and S. Kalimantan, Sabah, Sarawak).
Habitat & Ecology An early successional tree in lowland forest in Borneo. This species occurs in large gaps within primary forest, in logged forest, and is often found early in succession following shifting agriculture. It appears to have a broad tolerance of variation in soil moisture levels, as it occurs on hills and ridges as well as along streams and in wetter areas. Unlike the closely related M. griffithiana this species is not normally found in degraded swamp forests.
Notes 1. This species is closely related to M. griffithiana. It easily distinguished from that species in the deeply dissected (sometimes 5-lobed) leaves and deeply pectinate staminate bracteoles. See further discussion under that species.
2. Davies (1999) mentioned that M. motleyana in Borneo is restricted to eastern Sabah. Davies (2001) has recently found this species to be quite common in western Sarawak (see Davies 99017) although he has never seen it in northern Sarawak or Brunei. It is also yet to be collected from West Kalimantan.
Macaranga myriantha Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1008; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 324; Airy Shaw, Kew Bull. 37 (1982) 27; Whitmore, Gen. Macaranga Prodr. (2008) 188. Type: Teysmann s.n. (holo G-DC; photos K), Moluccas, Ambon, Haleke Ewan.
Macaranga robinsonii Merr., Philipp. J. Sci. 11, Bot. (1916) 284; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 184; Airy Shaw, Kew Bull. 37 (1982) 27.
Macaranga inermis auct. non Pax & K.Hoffm.: Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 148 quoad Kostermans 1216; Airy Shaw, Kew Bull. 37 (1982) 27, quoad specimina moluccensia.
Small trees to 10 m tall. Twigs slender, round, dark, glabrous. Stipules elliptic, to 5 mm long, blackish brown, thinly papery, glabrous but at first finely granular glandular, erect, soon caducous. Leaves: petioles to 5 cm, slender, round, swollen at both ends, glabrous or with spreading short pale hairs; blades elliptic, 1016 by 34.5 cm, thinly papery, brittle, base cuneate, with 2 basal glands at petiole insertion, margin entire, apex acuminate, drying dark brown, below with lower part of midrib usually with sparse spreading white hairs, and very sparse granular glands.Staminate inflorescences narrow racemes, to 4 cm long, 2 axis orders, glabrous; branches to 3 cm, divaricate, alternate, thread-like; peduncle 1 cm, main axis slender, terete, without axillary branches; bracts triangular, under 1 mm, subpersistent; flower clusters evenly spaced, c. 5 mm distant; bracteoles triangular, 1 mm across, apex acute, entire, persistent, glabrous, much smaller than flower cluster. Staminate flowers 712 per cluster, sessile; sepals free; stamens 7, anthers 4-locular. Infructescences racemes to 8 cm, axes flattened with sparse 2 cm spaced divaricate branches, lowermost opposite, axes slender, glabrous, brittle, bearing sparse fruits; bracteoles minute, ovate, under 1 mm, papery, glabrous persistent. Fruits round, 2 mm, thinly woody, granular glandular, sometimes with 14 short flat spines; pedicel 410 mm, slender; sepals elliptic, 1 mm, persistent; style basal, breaking off. Seeds unknown.
Distribution Moluccas (Morotai, Buru, Ambon).
Habitat & Ecology Forest, 9001000 m.
Macaranga neobritannica Airy Shaw, Kew Bull. 25 (1971) 531; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 151; Gen. Macaranga Prodr. (2008) 190. Type: Sayers NGF 21926 (holo K; iso SING), Papua New Guinea, New Britain Distr., Kandrian Subdistr., Pirilongi.
Small treea, to 8m. Twigs fairly stout, 5 mm, weakly angled, finely pubescent, becoming glabrous, sometimes with stipule scars. Stipules oblong-ovate, 1013 by 35 mm, apex acuminate, chestnut brown, papery, at first both sides tomentose, erect, subpersistent. Leaves: petioles 410 cm, slender, 2 mm diam., terete or slightly flattened, glabrous or with sparse long hairs, weakly kneed at top; blades variable, elliptic, 24 by 8 cm, to ovate, 1523 by 813 cm, papery, base usually broadly rounded and minutely cordulate or truncate, occasionally very broadly very shallowly cordate, rarely cuneate, with several inconspicuous small glands near petiole insertion, margin sometimes crenate or repand, with tiny inconspicuous distant glandular teeth, apex caudate, 23 cm, usually drying grey green, glabrous except sparsely hairy near petiole below, closely granular glandular below, 3 main and several very minor nerves from base. Staminate inflorescences delicate panicles to 10 cm, with 3 axis orders, slightly hairy or glabrous; branches arising from flower clusters, mostly ascending; peduncle 24 cm, slightly angled, lowest branches opposite, with axillary branches; bracts oblong, to 10 by 2 mm, apex acuminate, dark brown, papery, erect, usually persistent; secondary and tertiary branches sinuous thread-like; flower clusters evenly up to 5 mm spaced; bracteoles variable, narrowly elliptic, some 5 mm, apex acute, but apical ones smaller, ovate, to 2.5 by 2mm, apex obtuse, fleshy, with adaxial patellar glands, entire, glabrous, spreading, persistent, smaller than cluster. Staminate flowers c.10 per cluster (fewer at tips), sessile; stamens 2, anthers 4-locular. Infructescences spikes to 14 cm; axis flattened, 1.5 mm diam., glabrous or sparsely hairy, fruits usually just as a terminal cluster of 13, rarely 1 or 2 lateral clusters too, subtended at first by stalked elliptic leafy dentate papery bracts to 10 by 2 mm. Fruits when young enclosed in calyx, globose or shallowly dicoccous, 5(10) by 9(15) mm, thinly woody, densely covered with soft hairy almost terete 3 mm somewhat caducous spines; pedicel stout, 2 mm pubescent; calyx persistent; style apical, 2 mm, plumose, caducous. Seeds round, 4 mm, irregularly ridged, with sarcotesta.
Distribution East New Guinea, Bismarck Archipelago (New Britain, New Ireland), Admiralty Islands (Manus).
Habitat & Ecology Primary or disturbed forest. Sometimes on limestone. Alt.: sealevel to 1050(2000) m.
Notes 1. A polymorphic species, the variants united by intermediates. The form mentioned by Airy Shaw falls within the range of the species as described here.
2. Kuduk & Naoni UPNG 6196 from 2000 m on Mt. Ehuhu near Mt. Lamington differs in having young fruits totally spineless, globose, 5 mm diam.
3. Frodin & Morren 3421 is provisionally placed here. It differs in having very long stipules, 40 by 5 mm, and the truncate leaf base minutely auriculate at the petiole.
3. Frodin LAE 52368 from Manus is a staminate collection with very young flowers. The inflorescence bracts are thick, dentate and velvety and the inflorescences are borne mainly behind the leaves. It could represent a new species.
Macaranga noblei Elmer, Leafl. Philipp. Bot. 2 (1910) 678; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 317; Merr., Enum. Philipp. Flow. Pl. (1923) 442; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 35; Whitmore, Gen. Macaranga Prodr. (2008) 193. Type: Elmer 10816 (holo PNH, lost; iso K), Philippines, Mindanao, Davao Dist., Todaya (Mt. Apo).
Trees 8 m. Twigs fairly slender, 8 mm diam., angled, finely tomentose. Stipules not seen. Leaves: petioles 15 cm, fairly slender, round, weakly striate, at first finely velvety; blades deltoid, 1630 by 1528 cm, thinly leathery, base shallowly to 15 mm peltate, very broadly truncate with inconspicuous glands near petiole insertion, margin with distant inconspicuous glandular teeth, apex caudate, 3 cm, usually drying dark brown, below reticulations conspicuous, slightly raised, velvety to touch, main nerves with short white tufted hairs, granular glandular, above with very fine tufted evanescent short white hairs. Staminate unknown. Pistillate inflorescences 35 cm panicles, 3 orders of divaricate axes, furfuraceous, round; lowest branches opposite; flower clusters evenly spaced with 24 flowers; bracts caducous; bracteoles roundish, 2 mm, deeply regularly bluntly toothed, furfuraceous, caducous. Fruits strongly dicoccous, 5 mm broad by 3 mm long, smooth, thinly woody with granular glands; pedicel slender, 45 mm; calyx persistent; stigmas small, apical, rugose. Seeds 2, round, coarsely warty.
Distribution Philippines (Leyte, Surigao, Mindanao).
Note Staminate parts unknown, but a staminate tree from Luzon, Isabela, Sierra Madre mountains (Gutierrez 78248, K!) is close to M. noblei, conspicously differing in relatively longer leaves (23 by 16 cm), the base very shallowly broadly cordate, only 5 mm peltate and without raised reticulations below, and only a few of the hairs tufted. Stipules caducous, elliptic, 40 by 7 mm, widest at base, long acuminate, dark brown, papery, densely white-tomentose. Staminate inflorescences 20 by 8 cm panicles, 3 orders of divaricate axes, furfuraceous, somewhat flattened, main axes striate, with occasional axillary branches, ultimate branches thread-like with 8 mm spaced flower clusters; bracts ovate, 1 mm, persistent; bracteoles roundish, 2 mm, persistent, deeply regularly bluntly toothed, furfuraceous. Stamiante flowers 24 per cluster, pedicellate, tiny, 0.3 mm; calyx divided (?); stamens 2, anthers 4-locular. This tree is like a hairy form of M. amplifolia. More collections are needed to resolve the relationship of these two entities.
Macaranga novoguineensis J.J.Sm., Nova Guinea Bot. 8 (1912) 789, t. 138; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 363 (nova-guineensis); J. Arnold Arb. 34 (1953) 246; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 151; Gen. Macaranga Prodr. (2008) 193. Type: von Rφmer 514 (holo L), Indonesia, Papua, Noordfluss im Urwalde der Ebene.
var. novoguineensis
var. novoguineensis: Whitmore, Gen. Macaranga Prodr. (2008) 193. Type: as the species.
Shrubs or small trees, 3(8) m tall. Twigs slender, 23 mm diam., weakly angled, densely usually flavous pubescent. Stipules (narrowly) elliptic, 10 by 1 mm, slightly curved, pubescent, erect, subpersistent. Leaves: petioles 36 cm, slender to fairly stout, 24 mm diam., terete, pubescence as twigs, kneed at top; blades ovate to broadly ovate, 1216 by 712 cm, thickly papery, base broadly rounded (rarely broadly cuneate), at petiole sometimes very shallowly cordate or minutely peltate, with several small inconspicuous glands, apex acute or sometimes abruptly acuminate, drying khaki or dark brown, above lower midrib tomentose with white stiff solitary and tufted hairs, lamina sometimes with scattered solitary hairs, below dark granular glandular and densely to sparsely tomentose on midrib and nerves, hairs mostly solitary a few tufted, basal secondary nerves inserted at petiole. Staminate inflorescences open sparsely branched panicles, 18 by 6 cm, with 3 axis orders, densely fulvous tomentose, branches divaricate to 450; peduncle 1520 mm; main axis slender, flattened, 1.5 mm across; secondary branches to 7 cm, sometimes opposite, without accessories; bracts ovate, 1 mm, apex acute, persistent, tomentose; secondary branches slender; flower clusters contiguous in catkin-like clusters to 510 mm long at tips of secondary branches and as 5 mm tertiary branches; bracteoles oblong, 3 by 2 mm, apex acute, slightly cucullate, upper part with large regular gland-tipped teeth, pinnately nerved, pubescent without, persistent, glabrous within, (reminiscent of Drynaria frond), longer than flower cluster. Staminate flowers in clusters of 6, shortly pedicellate; sepals free; stamens 3, anthers 4-locular. Infructescences (4)10 cm spikes, with clusters of 2 or 3 terminal fruits, subtended by caducous leafy elliptic 15 by 4 mm bracts, granular glandular, tomentose on nerves, axis 1.5 mm diam., terete to slightly flattened, densely flavous pubescent. Fruits mostly strongly bilocular, loculi globose, 6m m diam., thinly woody, closely beset with soft slender hairy 6 mm spines, usually wine red (and drying reddish); pedicel stout, 1 mm or less, pubescent; calyx tiny, persistent; stigma 1 mm, stout, papillose. Seeds somewhat lenticular (? immature), 5 mm diam., coarsely verrucose, with sarcotesta.
Distribution Moluccas (Aru) and throughout New Guinea.
Habitat & Ecology Lowlands primary alluvial and ridge forest and secondary forest; 150(500) m.
Notes Smith (1912) said this has some androgynous inflorescences; cf. M. glaberrima var. schoddei.
2. The Aru collection (van Balgooy 6685) has the leaves broadest near the base.
3. Three collections have green fruits (Amiaba River south of Madang: NGF 42886, NGF 45852; Sepik: Ledermann 6964a). The leaf granular glands are not dark in NGF 45852.
var. glabra Whitmore
var. glabra Whitmore, Gen. Macaranga Prodr. (2008) 194. Type: Stevens LAE 58496 (holo K), Papua new Guinea, West New Britain, crater rim of North Son (extinct).
Differs from the typical variety in elliptic leaf blaes and complete absence of hairs.
Distribution East New Guinea (Morobe and Central Dist.) and Bismarck Archipelago (New Britain).
Habitat & Ecology Primary lower montane forest, 500990 m.
Note This lower montane, glabrous, narrower leaved variety is distinctive. The only staminate specimen seen does not have the flower clusters crowded.
Macaranga nusatenggarensis Whitmore, Gen. Macaranga Prodr. (2008) 194. Type: Schmutz 4428 (holo K; iso L), Indonesia, Nusa Tenggara, Flores, Manggerai.
Twigs slender, 3 mm diam. angular, at first finely golden furfuraceous, soon glabrous. Stipules ovate-elliptic, to 14 by 4 mm, apex long acuminate, occasionally caudate, mid to dark brown, papery, finely golden pubescent, mostly erect, persistent, crowded at twig tip. Leaves: petioles 810 cm, slender, round, striate, golden furfuraceous to subglabrous; blades round to deltoid, 1116 by 711 cm, papery, base broadly rounded or less commonly abruptly truncate, 23 cm peltate, margin with small distant glandular teeth, apex acuminate, drying khaki brown at least below, below closely golden granular glandular and sparsely pubescent on main nerves, spiders-web nervation. Staminate inflorescences narrow axillary panicles, to 14 by 4 cm, with 4 orders of axes; main axis slender, flattened, striate, weakly furfuraceous; basal peduncle to 8 cm; secondaries golden furfuraceous, spreading, to 3 cm, growing out of a flower cluster, alternate, lower few with axillaries, tertiaries to 1 cm, with crowded quaternaries to 5 mm, bearing crowded flower clusters with 56(14) flowers; bracts subtending all axes, ovate, to 5 by 4 mm, apex acute, glabrous, entire, persistent; bracteoles shorter than flower clusters, ovate, to 2 by 2 mm, apex acute, often with a few irregular coarse teeth, thick, granular glandular on both surfaces. Staminate flowers c. 15 per cluster, globular, 0.5 mm; pedicel 0.5 mm; sepals free; stamens 4, anthers 3-locular. Infructescences 7 cm, main axis slender, flattened, striate, weakly furfuraceous, with a few 2 cm branches near apex, bearing 13 fruits subtended by persistent leafy bracts, ovate, 20 by 8 mm, apex acuminate, deeply coarsely elongate- toothed, papery, glabrous or weakly furfuraceous, conspicuously nerved. Fruits sessile, depressed globose, 13 lobed, 5 by 10 mm, with sparse, mainly apical, 10 mm, sinuous spines, closely golden granular glandular, seated on small persistent calyx. Seeds round, coarsely knobbly, with sarcotesta.
Distribution Lesser Sunda Islands (Bali, Flores, Sumba, Sumbawa, Timor) and Moluccas (Tanimbar).
Notes 1. Resembles M. tanarius var. tanarius, differing mainly in the narrower stipules, to 14 by 4 mm, staminate inflorescences with 4 not 3 orders of axis, crowded flower clusters and bracteoles small, 2 by 2 mm, shorter than flower clusters, coarsely irregularly toothed.
2. Schmutz 3358 has leaves glaucous below, reminiscent of M. parabicolor, and with revolute margins.
Macaranga ovatifolia Merr., Philipp. J. Sc. 16 (1920) 562; Enum. Philipp. Flow. Pl. (1923) 442; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 35; Whitmore, Gen. Macaranga Prodr. (2008) 198. Type: Ramos & Pasgasio BS 34683 (holo PNH, lost; iso K), Philippines, Mindanao, Surigao Prov.
Small tree. Twigs stout, 6 mm diam., rounded, with conspicuous leaf scars, glabrous. Stipules elliptic, 15 by 3 mm, erect, blackish, papery, glabrous, caducous. Leaves: petioles 17 cm, slender, round, slightly striate, glabrous; blades ovate, to 20 by 17 cm, thinly leathery, base broadly shallowly cordate, margin entire, apex acute, brown, glabrous, with very fine dark granular glands below. Stamiante unknown. Infructescences behind leaves, narrow diffuse 8 cm panicles, with 2 somewhat flattened branching orders; branches divaricate, slender, sinuous, sparsely furfuraceous; bracts tiny, ovate, 1 mm, persistent; bracteoles ovate, 1 mm, coarsely shallowly dentate, persistent, furfuraceous; clusters a few mm apart. Fruits in groups of 3, monococcous, round or slightly broader than long, 3 mm, thinly woody, smooth with sparse granular glands; pedicel under 1 mm; calyx persistent. Seeds round, finely verrucose, wholly enclosed in a thin sarcotesta.
Distribution Philippines (known from the type only).
Macaranga pachyphylla Mόll.Arg. in DC., Prodr. 15, 2 (1866) 999; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 319; Whitmore in Airy Shaw, Kew Bull. 36 (1981) 321; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 375; Whitmore, Gen. Macaranga Prodr. (2008) 199. Type: Zollinger 18152 (holo GDC; iso L), Sumatra, Lampong.
Macaranga curtisii Hook.f., Fl. Brit. India 5 (1887) 448; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 318; Ridl., Fl. Malay. Penins. 3 (1924) 301; Whitmore, Tree Fl. Malaya 2 (1973) 107; in Airy Shaw, Kew Bull. 36 (1981) 319; Airy Shaw Kew Bull. 26 (1972) 287.
Macaranga semiglobosa J.J.Sm., Meded. Dep. Landbouw. 10 (Bijdr. 12) (1910) 503; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 320; Backer & Bakh.f., Fl. Java 1 (1963) 488; Whitmore in Airy Shaw, Kew Bull. 36 (1981) 319. Lectotype (designated by Whitmore, 2008): Koorders 30545ί (holo BO).
var. pachyphylla
var. pachyphylla: Whitmore, Gen. Macaranga Prodr. (2008) 199. Type: as the species.
Small trees to 8 m. Twigs slender, to 5 mm diam., terete, with conspicuous encircling stipule scars, softly fawn tomentose to glabrous and glaucous. Stipules ovate, 12 by 10 mm, apex broadly acute, dark brown, papery, glabrous to tomentose in centre, with sparse granular glands, erect becoming recurved, subpersistent. Leaves: petioles 817 cm, slender, 2 mm diam., terete, weakly to densely tomentose with at least a few spreading hairs; blades ovate, 918(30) by 613(25) cm, coriaceous, base broadly rounded, 28 cm peltate, margin sometimes recurved with protruding glandular teeth, apex acute to acuminate for 1 cm, drying chocolate brown, below with sparse granular glands and velvety from spreading hairs on all main nerves, sometimes slightly glaucous, 5 main nerves at petiole insertion. Staminate inflorescences crowded, panicles dense, 9 by 3 cm, becoming open, 19 by 7 cm, 3 axis orders, tawny tomentose to glabrous branches ascending to spreading, to 7 cm long; peduncle 14 cm; main axis slender, 1.53 mm diam., terete or flattened, coarsely ridged; lower branches often opposite, sometimes alternate, sometimes with a few axillary branches; bracts variable, ovate, 310 by 15 mm, apex obtuse-acute or acuminate, dark brown, papery, both sides slightly rusty pubescent to subglabrous, soon caducous; ultimate branches slender, bearing crowded flower clusters; bracteoles variable, erect, overlapping, persistent, ovate, strongly concave, 14 by 13 mm, usually with abrupt 1 mm caudate tip, margin entire, both sides densely rusty furfuraceous. Staminate flowers 46, 0.5 mm diam.; pedicel 0.5 mm;, sepals hairy; stamens 34, anthers 4-locular. Infructescences 410 cm spikes, with a few fruits towards apex; axis terete or flattened, tomentose to glabrous. Fruits globose, often slightly depressed, to 8 mm diam., black, leathery, shallowly grooved, closely granular glandular except grooves; pedicel 10 mm, stout, tomentose; sepals large (1 mm), persistent, papery; stigmas small, 1 mm, persistent, broad, apical. Seeds globose, striate, with sarcotesta.
Distribution Thailand, Malay Peninsula, Sumatra, Java.
Habitat & Ecology Primary mountain rain forest, (100)5001500 m.
Notes 1. The type from Lampung, Zollinger 1815Z (L!), has unusually large leaves and very young inflorescences, but there is no doubt it is the species widely known till now as M. curtisii.
2. A rather polymorphic species. Most gatherings have conspicuous hairy twigs and velvety leaves but intermediates to the virtually glabrous and highly distinctive var. glabra occur. Those with the twigs glabrous and often glaucous, but hairy leaves I place here, restricting to var. glabra the glabrous gatherings.
3. Sometimes with a superficial resemblance to M. tanarius var. tomentosa which, however, has staminate bracteoles dentate and fruits spiny.
4. Iwatsuka et al. S 456 differs in possessing very large pyriform fruits, 15 mm long.
5. I have identified Koorders 23483β, and 30545β as syntypes of M. semiglobosa. The BO sheets are determined in Smith's hand. Of these 23483β from Pasuruan, Malang, is the glabrous variety, therefore I select Koorders 30545β as lectotype of M. semiglobosa.
var. glabra (Whitmore) Whitmore
var. glabra (Whitmore) Whitmore, Gen. Macaranga Prodr. (2008) 200. Macaranga curtisii Hook.f. var. glabra Whitmore, Kew Bull. 25 (1971) 240; Tree Fl. Malaya 2 (1973) 111; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 151, in Airy Shaw, Kew Bull. 36 (1981) 319. Type: Whitmore KEP FRI 312 (holo Kl iso L), Peninsular Malaysia, Selangor, Gunung Bunga Bua.
Macaranga brooksii Ridl., Kew Bull. (1925) 90; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 151 passim; in Airy Shaw, Kew Bull. 36 (1981) 319.
Macaranga aff. puncticulata sec.Corner, Wayside Trees of Malaya (1940) 269.
Macaranga aff. puncticulata (montane form) sec. Whitmore, Malay. Nat. J. 20 (1967) 98.
Differs from var. pachyphylla in: Twigs glabrous sometimes glaucous. Stipules very soon caducous, almost glabrous. Leaves: petioles glabrous or with a few scattered hairs, sometimes slightly glaucous; blades below closely granular glandular, glabrous or with (usually scattered) hairs on (main) nerves. Inflorescence axes and fruiting pedicels furfuraceous.
Distribution Malay Peninsula, Sumatra, Java, Borneo (Sabah, Brunei, Sarawak).
Habitat & Ecology Primary mountain forests, 5001500 m.
Note This glabrous or nearly glabrous variety is partially sympatric with the type variety whose habitat it shares.
Macaranga palustris Whitmore, Kew Bull. 34 (1979) 604; in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 151; Gen. Macaranga Prodr. (2008) 200. Type: Foreman & Katik LAE 52014 (holo K; iso L, SING), Papua New Guinea, Western Dist., Kiunga Subdist., W of Kiunga Airstrip.
Small trees 9 m. Twigs stout, 4 mm diam., angled, becoming coarsely striate, with conspicuous leaf scars, densely tomentose, golden at first. Stipules oblong, 7 by 3 mm, apex acute, chestnut brown, papery, outside patchily tomentose, erect, long-persistent. Leaves: petioles 67 cm, stout, 3 mm diam., terete, densely tomentose, weakly kneed at both ends; blades ovate-elliptic, c. 19 by 8 cm, leathery, base rounded, cordulate with overlapping auricles and numerous small glands near petiole insertion, margin strongly sinuate, slightly recurved, with tiny protruding glandular teeth, apex shortly acuminate, drying grey above, pubescent on midrib and secondaries above and on all nerves below, sparsely dark granular glandular below, reticulations raised below, irregularly bullate between scalariform tertiaries, secondary nerves strictly pinnate. Staminate unknown. Infructescences behind leaves, open panicles to 18 by 7 cm, with 3 axis orders; main branches to 4 cm, spreading or ascending; peduncle to 3 cm; main axis angled, 2 mm diam., finely furfuraceous; branches alternate; bracts mostly persistent, small, triangular, 1 mm, apex acute, thick, furfuraceous, ultimate branches to 1 cm, stout. Fruits solitary or paired, 5 mm diam. (? young), globose or sometimes bilobed, wall woody, densely black granular glandular, at first rusty pubescent, with a few slightly raised 1 mm smooth bosses; pedicel stout, 2 mm; calyx furfuraceous, persistent; styles 1 or 2 short, apical, caducous, smooth. Seeds globose, coarsely shallowly verrucose, with sarcotesta.
Distribution New Guinea (known from type only).
Habitat & Ecology Lowland swamp forest, 20 m.
Notes 1. Unusual for a New Guinea species to be found in a swamp habitat.
2. Reminiscent of the Conifera species group of the Sunda Shelf. Macaranga palustris and M. hengkyana are unique in the group in their strictly penninerved leaves and lowland swamp habitat.
Macaranga papuana (J.J.Sm.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 368; L.M.Perry, J. Arnold Arb. 34 (1953) 206; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 152; Gen. Macaranga Prodr. (2008) 201. Macaranga hispida (Blume) Mόll.Arg. var. papuana J.J.Sm., Nova Guinea Bot. 8 (1910) 234. Type: Jaheri 234 (holo L), Indonesia, Papua, Merauke Dist., Sungai Muroka.
Macaranga ovalifolia Ridl., Trans. Linn. Soc. Bot. 9 (1916) 148; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xvii add. vii (1924) 186.
var. papuana
var. papuana: Whitmore, Gen. Macaranga Prodr. (2008) 201. Type: as the species
Trees to 18 m. Twigs slender, angular, 3 mm diam., at first furfuraceous, with conspicuous stipule scars. Stipules united, apparently single, forming an erect slightly swollen 23.5 cm by 3 mm terminal bud, finely ochre tomentose, papery, caducous. Leaves: petioles c. 7 cm, terete, 2 mm diam., ochre furfurescent; blades ovate-elliptic, 921 by 4.514 cm, papery, base cuneate to truncate (rarely very shallowly broadly cordate) with 2 small gland bearing auricles at petiole insertion, margin repand, apex acuminate, drying khaki brown, above glabrous (except midrib puberulous), below nerves furfuraceous with close minute clustered hairs and black granular glandular. Staminate inflorescences sparsely branched delicate sinuous racemes to 20 cm; main axis slightly flattened, 1.5 mm diam., ochre furfuraceous; peduncle to 7 cm; branches long, lowest opposite, usually with axillary branches, steeply ascending, thread-like with evenly spaced c. 3 mm distant tiny flower clusters; bracts soon caducous, shortly stalked, elliptic, to 1 cm, bearing numerous small adaxial patellar glands; bracteoles ovate, 1 mm across, apex obtuse, cucullate, margin crenate, papery, sparsely long-hairy outside, persistent, with a caducous suborbicular central lobe bearing patelliform glands, shorter than cluster of c. 15 flowers. Staminate flowers tiny, shortly pedicellate; stamens 45, stout, anthers 3-locular. Infructescences spikes to 6 cm, with a few near-terminal solitary fruits;, axis flattened, 1 mm across, ochre furfuraceous; bracts elliptic, stipule-like, 1 cm, erect. Fruits weakly bilobed, 8 by 10 mm (immature), densely covered with 3 mm slightly flattened, soft, sinuous, tousled, hairy spines, granular glandular. Seeds unknown.
Distribution New Guinea.
Habitat & Ecology Primary lowland forest.
Notes 1. This species is distinctive in the gland-bearing auricles at the leaf base. The clustered hairs on the leaf below are reminiscent of M. hispida.
2. Lam 1502 differs in having the leaves leathery, weakly bullate, the nerves all raised below, and in its occurrence in the mountains at 1420 m near Doormantop.
var. glabristipulata Whitmore
var. glabristipulata Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 152; Gen. Macaranga Prodr. (2008) 202. Type: Streimann LAE 51849 (holo K; iso L), Papua New Guinea, Western Dist., Kiunga.
Differs from the type variety in being entirely glabrous and with the leaves often slightly glaucous below.
Distribution New Guinea (Mt. Carstensz., Western Dist.).
Habitat & Ecology Lowlands, regrowth forest.
Macaranga parabicolor Whitmore, Gen. Macaranga Prodr. (2008) 202. Type: Schmutz 3880 (holo K; iso L), Indonesia, Nusa Tenggara, Flores, west at Nunang.
Small tree to 20 m. Twigs stout, to 1 cm diam., round, pruinose and glabrous or rusty furfuraceous, with stipule scars. Stipules ovate, 20 by 10 mm, apex acuminate, fawn furfuraceous, papery, erect, soon caducous. Leaves: petioles 1013(40) cm, round or angled, 3 mm diam., striate, pruinose or rusty furfuraceous; blades very broadly ovate, 1845 by 1740 cm, thinly leathery, base very broadly rounded to subtruncate, deeply 512 cm peltate, margin with tiny distant protruding glands, apex acuminate, drying dark brown above, mid brown below, nerves and surface finely densely pubescent below, velvety to the touch, close darkish granular glands, main nerves palmate, tertiaries and quaternaries scalariform. Staminate inflorescences open panicles, 16 by 6 cm, with 3 orders of stout axes; latter angular, striate, flattened, tawny furfuraceous, secondaries to 11 cm, lower ones opposite, a few 1 cm tertiaries, branches developing through flower clusters; bracts and bracteoles round, to 5 by 4mm, entire, apex caudate, 1 mm, persistent, patent or reflexed, both sides furfuraceous, 2 patellar glands developing to cover adaxial surface, longer than the conspicuous bun-like, 4 by 3 mm crowded flower clusters. Staminate flowers c. 30 per cluster; pedicel 0.5 mm; calyx 1 by 0.5 mm, pubescent; stamens 5, anthers 3-locular. Infructescences narrow racemes to 7 cm long; branches 2 cm, opposite; basal peduncle 24 cm long, 2 mm diam.; axes stout somewhat flattened, striate, some times pruinose, weakly golden furfuraceous; bracts and bracteoles caducous, not seen. Fruits depressed roundish, to 5 by 6 mm, shallowly 4-lobed, weakly shouldered, the lobes very weakly grooved or not, single and also crowded towards apex, drying blackish, wall leathery, golden furfuraceous becoming subglabrous, smooth, closely granular glandular except at base of sutures between lobes; pedicel stout, 3 mm, golden furfuraceous; calyx 2 mm across, golden furfuraceou, persistent; styles free, on top of lobes, more or less erect, 2 mm. Seeds with red sarcotesta, round, surface shallowly sculptured.
Distribution Lesser Sunda Islands (Flores, Sumbawa), Moluccas (Kai Islands).
Habitat & Ecology Mountains, 1001200 m.
Note With similar leaves to those of M. bicolor of Philippines, but amply distinct in the more robust staminate inflorescences with lower branches opposite, staminate bracteoles with patellar glands, longer 7 cm infructescences, short 3 mm fruiting pedicel, and larger 2 mm stigmas.
Macaranga pearsonii Merr., Philipp. Journ. Science 29 (1926) 383; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 153; S.J.Davies, Harvard Pap. Bot. 6 (2001) 431. Type: Castro 1672 (holotype: UC, not seen; isotypes: A, K), Malaysia, Sabah, Banguey Island.
Large tree to 35 m tall and 50(60) cm dbh, often smaller in open areas; twigs 612 mm in diam., glabrous, usually densely glaucous, hollow, housing ants. Bark pale whitish-grey to light brown, smooth, hoop marked. Stipules broadly ovate, 914 by 913 mm, erect to saccate, glabrous, the pair not completely encircling the twig, coriaceous, producing food-bodies on the upper surface, up to 3 pairs present on the shoots. Leaves: petioles terete, slender, 1030 cm long, glaucous towards the base, finely pubescent towards the apex; blades ovate to very broadly ovate, 1328 by 1330 cm, deeply trilobed, dissected to c. 1/2 of the leaf length rarely somewhat less, lateral lobes 1/2 to 4/5 the size of the central lobe, not peltate or sometimes to 10 mm peltate, base broadly rounded, cordate, margin entire, usually with 16 prominent conical nectaries at the leaf base adjacent to the petiole insertion, apices shortly acute, adaxial surface glabrous or scattered with minute ferrugineus hairs along the veins near the petiole insertion, abaxial surface densely minutely furfuraceous tomentose on the veins and scattered with long erect silvery hairs and densely glandular punctate; 1° venation palmate with 5 prominent veins, 59 subprominent veins running from the petiole insertion to the basal leaf margin, 2° venation scalariform terminating at the margins in prominent conical nectaries or looping just before the leaf margins, 3°4° venation densely scalariform, finest nerves densely reticulate; young leaves reddish brown, upper surface densely covered in fine ferrugineus hairs, lower surface with dense erect silvery hairs. Staminate inflorescences paniculate, erect, 1540 by 1025 cm, all axes densely covered in erect silvery hairs, the basal axis sometimes ± glabrous, to 4 axis orders, main axis flattened, first pair of secondary branches ±opposite without accessory branches; bracts narrowly ovate, 510 by 36 mm, margin entire, apex acute, glabrous or with scattered silvery hairs on the margins, persistent; flower clusters with c. 10 flowers, spially arranged on inflorescence branches; bracteoles ovate-elliptic, 36 by 24 mm, enclosing flower clusters, margin deeply irregularly dissected, apex acute to acuminate, veins slightly raised on the abaxial surface, both surfaces densely covered with fine silvery and reddish brown hairs. Staminate flowers c. 1 mm long, pedicellate; sepals free, apex densely covered in minute red-brown hairs; stamens 23; anthers 4-locular. Pistillate inflorescences paniculate, erect, 825 by 816 cm, densely branched, finely silvery pubescent throughout, up to 3 axis orders; bracts ovate, 510 mm long, margin entire, glabrous or with scattered hairs on the margins, caducous. Pistillate flowers c. 1.5 mm long, solitary in bract axils; calyx urceolate, c. 1 mm long, scattered with fine silvery and ferrugineus hairs, persistent; ovary 2-carpellate, c. 1 mm long; styles c. 1 mm long, free, persistent; stigma not dissected. Fruits 34 by 56 mm, bilocular, locules globose, pedicellate, evenly covered in sticky glandular exudate, without processes; pedicel 47 mm long, densely pubescent. Seeds lenticular, c. 2.5 mm in diam., black, with shallow grooves, encased in a fleshy pale purple aril.
Distribution Borneo (East and South Kalimantan, Sabah). This species is known from Sabah east of Ginning Kinabalu and the Crocker Range, and East and South Kalimantan.
Habitat & Ecology A very common early successional tree in lowland dipterocarp forest in east Borneo to c. 700 m elevation. Macaranga pearsonii grows to a large size in intact forest.
Note The densely glaucous stems, hairy inflorescence axes, and prominent conical nectaries on the margins of the leaf bases distinguish this species from its close relatives. It is probably most closely related to M. hosei and M. rufescens. It differs from these species most obviously in having cupulate or saccate stipules only a few of which remain persistent on the twigs.
Macaranga pentaloba S.J.Davies, Harvard Pap. Bot. 6 (2001) 433. Type: Davies 2000151 (holotype: A), Malaysia, Sarawak, near Pare Bata camp, from Ba'Kalalan to Gunung Murud, Kelabit Highlands.
Locally common submontane pioneer tree to 20 m tall and 30 cm dbh; twigs stout, 1830 mm in diam., terete though strongly ribbed when dry, densely covered in 23 mm long simple erect hyaline hairs, the hairs falling as the stems mature, tufted hairs absent, moderately to densely glaucous, exuding copious red latex when cut. Stipules huge, broadly ovate to oblong, to 13 by 7 cm, smaller in mature plants, erect, apex acute to rounded, drying black and chartaceous, sparsely covered in 23 mm long simple erect hyaline hairs, persistent in a cluster at the shoot apex. Leaves: petioles 3050 cm long, stout, to 9 mm in diam., terete or slightly flattened on the adaxial surface near the base, sparsely to densely covered with simple very long erect hyaline hairs, slightly glaucous; blades huge, broadly ovate, 4070 by 3060 cm, probably even larger in saplings, thickly coriaceous, 57 lobed, dissected to c. 1/3 with ovate to broadly ovate lobes, deeply peltate to 512 cm, base broadly rounded to truncate, margin slightly dentate, pubescent, apices acute, adaxial surface sparsely to densely covered in very long erect soft hyaline hairs and without tufted hairs, abaxial surface densely pubescent with long simple soft hyaline hairs and densely punctate glandular; 1° venation palmate with c. 8 prominent veins, 2° venation scalariform looping and terminating at the margins in conical nectaries, 3°4° venation densely scalariform, finest nerves densely reticulate and prominent; leaf; leaf; leaf; young leaves not seen. Flowers and fruits not seen.
Distribution Borneo (Sarawak). Davies (2001) recorded this species from only two locations: near Bario and from Gunung Murud. It will be interesting to see how much more widely this species is distributed.
Habitat & Ecology Macaranga pentaloba is a striking new species recently discovered from submontane forest in the Kelabit Highlands. It appears to be similar to the closely related M. gigantea in being a relatively large early successional tree. Following logging this species has abundant regeneration.
Note Macaranga pentaloba is closely related to M. gigantea. Both species have enormous lobed leaves, stout twigs, and large coriaceous stipules that dry black and papery and persist at the shoot apex. Macaranga pentaloba differs from M. gigantea in having extraordinary 57 lobed leaves; M. gigantea almost always has trilobed leaves. Macaranga pentaloba also has distinctive long erect soft hyaline hairs on upper and lower leaf surfaces, twigs and stipules, and does not have tufted hairs which are common in M. gigantea. The young stems of M. pentaloba are also moderately to densely glaucous, unlike those of M. gigantea.
Macaranga pepysiana Whitmore, Gen. Macaranga Prodr. (2008) 205. Type: Kerenga et al. LAE 77515 (holo K), Papua New Guinea, Admiralty Islands, Manus Prov., Lorengau, Mt. Dremsel.
Shrubs or small trees to 20 m. Twigs slender, terete, 2 mm diam., glabrous, ageing whitish. Stipules and bud scales subulate, 23 mm long, densely tomentose, erect, persistent. Leaves: petioles 1 cm, terete, 1 mm diam., glabrous, strongly kneed at both ends; blades elliptic, 1017.5 by 2.54.5 cm, papery, base long cuneate, without glands near petiole insertion, margin repand, apex gradually or abruptly tapering, blunt acuminate, drying chocolate brown, glabrous with or without sparse black granular glands below. Staminate unknown. Infructescences slender 4 cm spikes with axis thread-like (<1mm), angular, sparsely pubescent, bearing a few distant tiny 1 mm bracts and a single terminal fruit. Fruits weakly bilobed, 6 by 7 mm, thickly woody, densely beset with short stout sharp spines; pedicel 1 mm, pubescent; calyx persistent; styles 23, free, erect, apical, coarsely papillose, 1 cm, persistent. Seeds globose, 4 mm across, smooth, enclosed in an sarcotesta.
Distribution Admiralty Isldns (Manus Island).
Habitat & Ecology Primary lowland or lower montane rain forest, 300610 m.
Notes 1. The long apical styles are highly distinctive and reminiscent of a stack of poles.
2. An important part of Samuel Pepys' job as secretary to the Admiralty was to acquire the poles needed for ships' masts and spars.
Macaranga petanostyla Airy Shaw, Kew Bull. 25 (1971) 537; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 149; S.J.Davies, Harvard Pap. Bot. 6 (2001) 415. Type: Chew 1612 (holotype: K; isotypes: L, SAN, SING), Malaysia, Sabah, Mamut River, Gunung Kinabalu.
Small submontane tree to 10 m tall with large to very large dark green glossy leaves; twigs stout, 712 mm in diam., mostly glabrous or the youngest twigs with c. 1 mm long sharply erect hyaline hairs, not glaucous, hollow, housing ants. Stipules ovate, 811 by 1012 mm, glabrous, recurved and appressed to the stem, succulent, producing food-bodies on the abaxial surface, up to 4 pairs present on the shoots. Leaves: petioles terete, quite stout, (10)1530 cm long, not glaucous, glabrous; blades glossy dark green, broadly to very broadly ovate, 2040(50) by 1540 cm, trilobed or tricusped and rarely some specimens with an additional pair of basal cusps, dissected to 1/81/4 of the leaf length, central leaf lobe up to 18 cm wide at the base and narrowing to a broadly acute apex, lateral lobes short and spreading with acute apices, 59 cm peltate, base very broadly rounded, margin entire, sometimes with small flattened protruding triangular nectaries, apices broadly acute, adaxial surface glabrous, abaxial surface mostly glabrous with minute silvery crisped hairs on the veins, not glaucous; 1° venation palmate with 810 prominent veins, 2° venation scalariform, looping near the leaf margins and ending in small flattened nectaries, 3°4° venation scalariform, minor veins densely reticulate; young leaves shiny crimson when fresh, densely and minutely ferrugineus pubescent. Staminate inflorescences paniculate, erect, 619 by c. 10 cm, basal unbranched rhachis very short 0.53 cm long, covered in erect sharp pointed c. 1 mm long hyaline hairs towards the base and becoming sparse to completely absent on distal axes, distal axes scattered with minute ferrugineus hairs, to 4 axis orders, main axis flattened, first pair of secondary branches opposite with accessory branches; bracts ovate-elliptic, 616 by 38 mm, margin entire, apex acute, mostly glabrous or scattered with fine ferrugineus hairs towards the base, caducous; flower clusters with 1520 flowers, spirally arranged and often so closely spaced on ulti mate inflorescence branches that they run into each other and appear almost spike-like; bracteoles elliptic to subulate, slightly broader at the base, 24 by 1.01.5 mm, erect, subtending the flower clusters but not enclosing the clusters, margin entire, apex caudate, scattered to densely covered with minute red-brown hairs. Staminate flowers c. 0.75 mm long, sessile; sepals fused, splitting irregularly to c. 1/5, apex densely covered in minute red-brown hairs; stamens 1; anthers 3-locular. Pistillate inflorescences paniculate, erect, 712 by 610 cm, stout, base of main rhachis densely covered in long sharply pointed hyaline hairs becoming sparse towards the apex, 2 axis orders, secondary branches ± opposite; bracts elliptic with caudate apex, glabrous or scattered with minute sparse ferrugineus hairs, some more distal bracts with 1 to several marginal conical nectaries, caducous. Pistillate flowers 34 mm long, solitary in bract axils; calyx urceolate with a truncate apex, c. 2 mm long, apex with fine ferrugineus hairs, persistent, splitting irregularly as ovary expands; ovary 45 carpellate, c. 3 mm long; styles 35 mm long with long and usually curled acuminate apices, fused at the base, free and distinctly spreading from c. 1/3 of the length, persisting to form a prominent 58 mm long crown at the fruit apex with the style apices remaining curled; stigma not dissected. Fruits (not quite mature) subglobose, compressed, 69 by 1013 mm, sessile, one discrete glandular patch on each carpel wall developing into a long slender horn-like process 46 mm long, covered in glandular, sticky exudate. Seeds subtriangular-ovoid, 44.5 mm in diam., black, with rather small coarse grooves and a small cruciform scar, encased in a fleshy aril (color not seen).
Distribution Borneo (Sabah). It is known from Gunung Kinabalu, the Crocker Range and Gunung Trus Madi.
Habitat & Ecology Macaranga petanostyla is a small large-leaved pioneer tree of submontane forest between 1000 and 1400 m. It occurs in logged forest, and along roadsides in primary forest. It is appears to be quite shade tolerant and occurs along rivers and in canopy openings in intact primary forest.
Note Macaranga petanostyla is quite closely related to M. calcicola which is endemic to forest over limestone in the Bau hills of western Sarawak near Kuching. Both of these species have glossy green leaves, erect hyaline hairs on the young twigs and basal inflorescence axes, staminate bracteoles that subtend but do not fully enclose the flower cluster, and adjacent flower clusters very close and almost continuous on the distal inflorescence axes. In addition to their strikingly different ecologies, M. petanostyla also differs from M. calcicola in having very long spreading styles which are fused for only c. 1/3 of the style length, much larger and usually more deeply dissected leaves, more glabrous leaves and twigs, and fruits with much longer horns. Macaranga calcicola also has one or several small conical nectaries on the margin of the staminate bracteoles; these appear to be absent in M. petanostyla.
Macaranga pilosula Airy Shaw, Kew Bull. 25 (1969) 105; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 152; Gen. Macaranga Prodr. (2008) 208. Type: Carr 15574 (holo K; iso L, SING), Papua New Guinea, Northern Prov., Isuarava.
Trees. Twigs 612 mm diam., slightly angular with close stipule scars, glaucescent, glabrous. Stipules ovate-oblong, 45 by 14 mm, more or less straight sided, tapering to acuminate tip, often with a prominent central nerve, dark brown with greyish blotches, papery, almost glabrous, glaucescent, erect, persistent. Leaves: petioles 1216 cm, terete, 3 mm diam., faintly ridged, glabrous with grey patches, or sparsely hairy, sometimes constricted at tip; blades ovate-oblong, 1928 by 1125 cm, papery or thinly leathery, base strongly rounded, 4(9) cm peltate, margins crenate to repand, sometimes with a 5 cm tip narrowing to a caudex, drying khaki brown, below sometimes glaucous, velvety with close spreading white hairs on all nerves (becoming almost glabrous), tiny inconspicuous golden granular or black glands sometimes only in patches, above at first densely hairy on main nerves elsewhere with sparse long white hairs, with spider's web nervation, tertiaries (faint above) scalariform. Staminate inflorescences narrow 16 cm racemes; main axis flattened, 3 mm across, glabrous near base, distally golden pubescent with grey patches; peduncle to 5 cm; branches arising from a flower cluster, ascending to 2 cm long, densely golden pubescent, a few short axillary branches; flower clusters crowded distally, bracteoles overlapping; bracts a few, small, triangular, 3 mm, apex acute, spreading tomentose; bracteoles roundish, 5 mm, apex shortly acuminate, strongly cucullate, parallel nerved, margin regularly, lightly pectinate, densely golden velvety on both surfaces, spreading, persistent, larger than flower cluster. Staminate flowers in clusters of 2530; stamens 7, anthers 4-locular. Infructescences sparsely branched racemes to 20 cm; main axis somewhat flattened, 3 mm across, glabrous or pubescent; peduncle to 13 cm; branches to 3 cm, subtended by bracts; latter ovate, 2 cm, apex acute, caducous. Fruits crowded at tips, subtended by subpersistent 2 cm stipule-like bracts, or dispersed and bracts caducous; fruits globose, 5 mm across (immature), golden pubescent, densely set with flattish 2 mm black spines; pedicel stout, 2 mm; calyx caducous. Seeds not seen.
Distribution West and southeast New Guinea, Bougainville.
Habitat & Ecology Lowlands and 1500 m.
Notes 1. Distinctive amongst its relatives in the big flat, erect stipules bearing a faint central nerve.
2. The West New Guinea collection (Polak 1333) has the fruits dispersed but otherwise is a close match.
Macaranga pleioneura Airy Shaw, Kew Bull. 23 (1969) 110; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 153; Gen. Macaranga Prodr. (2008) 209. Type: Brass 31192 (holo K; iso L), Papua New Guinea, Eastern Highlands Distr., Mt. Michael, W slopes.
Macaranga polyneura Gilli, Ann. Naturhist. Mus. Wien 83 (1980) 438.
var. pleioneura
var. pleioneura: Whitmore, Gen. Macaranga Prodr. (2008) 209. Type: as the species.
Shrubs or small trees, to 9 m tall; inovations sparsely puberulous. Twigs 3 mm diam., slightly angled, rarely rounded, occasionally striate, glabrous, blackish, slightly zigzag, internodes usually 1.5 cm or less. Stipules elliptic to narrowly elliptic, 810 mm long, apex acuminate, blackish, papery, outside densely tomentose, erect, soon caducous. Leaves: petioles c. 3 cm, variably flattened, 2 mm across, almost glabrous to densely shortly tomentose, not kneed; blades narrowly elliptic, 1117 by 47 cm, leathery, tapering gradually to cuneate base, with 2 large (to 4 mm) elongate marginal glands near petiole insertion, margin sometimes with tiny conspicuous protruding glands, slightly recurved and near base usually strongly recurved, apex acute to acuminate, drying grey green above, bright pale brown often yellowish below, occasionally coarsely bullate, glabrous, or slightly puberulous on main nerves below, densely golden granular glandular, secondary nerves close straight parallel, numerous (1222 pairs). Staminate inflorescences behind or crowded amongst leaves, open sparsely branched 10 cm long racemes, glabrous but with sparse granular glands; branches strongly divaricate, to 4 cm, arising from a flower cluster; peduncle 1 cm; main axis somewhat flattened, 11.2 mm across, striate; branches usually alternate, a few (sub)opposite; bracts elliptic, 7 by 1 mm, granular glandular, sometimes dentate with large marginal superficial glands, leathery, strongly recurved, soon caducous; flower clusters evenly, c. 3 mm apart; bracteoles ovate, 1.5 mm, apex blunt, fleshy, glabrous, erect, persistent, smaller than clusters of c. 4 flowers. Staminate flowers developing in sequence, shortly pedicellate; sepals thick, free, granular glandular; stamens c. 3040, anthers 4-locular. Pistillate flowers: sepals narrowly triangular, united at base, granular glandular. Infructescences amongst leaves, crowded spikes or sparsely branched racemes, to 10 cm long, as staminate but main axis flattened. Fruits in clusters of 28, tiny, 2 mm, globose, at first densely ochreous granular glandular, sometimes puberulous, with 06 weakly hairy 2.5 mm slender horns; pedicel long, slender, 15 mm, glabrous or puberulous; calyx persistent; style basal, densely plumose, 3 mm, caducous. Seeds globose, shallowly verrucous, with sarcotesta.
Distribution New Guinea (through the central mountains (mostly in east) and southeast Peninsula).
Habitat & Ecology Secondary montane forest, 9002400 m.
var. velutina Whitmore
var. velutina Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 153; Gen. Macaranga Prodr. (2008) 209. Type: Ridsdale & Woods NGF 33794 (holo K), Papua New Guinea, Cenral Prov., Goilala Subdistr., Woitape.
Macaranga hageniana Gilli, Ann. Naturhist. Mus. Wien 83 (1980) 437.
Differs in all parts persistently velutinous.
Distribution East New Guinea (Central, Southern & Western Highlands Provinces).
Macaranga pleiostemon Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv add. vi (1919) 24 (pleiostemona); L.M.Perry, J. Arnold Arb. 34 (1953) 255; Airy Shaw, Kew Bull. 23 (1969) 112; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)153; Gen. Macaranga Prodr. (2008) 210. Type: Ledermann 12493 (not seen), Papua New Guinea, Sepik Region, Felsspitze.
Macaranga pleiostemon Pax & K.Hoffm. forma pubescens L.M.Perry, J. Arnold Arb. 34 (1953) 256.
Shrubs or small trees, to 9 m. Twigs slender, 23 mm diam., angled, glabrous or finely furfurescent, dark brown. Stipules elliptic, to 8 mm, apex acuminate, papery, pubescent to subglabrous, erect, soon caducous. Leaves: petioles 12 cm, 1.5 mm diam., angled, glabrous, weakly kneed at top; blades elliptic to narrowly elliptic, 915(19) by 23(3.5) cm, thinly leathery, long-tapering to cuneate usually cordulate base, with 2 conspicuous elongate glands near petiole insertion, margin with tiny conspicuous protruding glands, usually slightly recurved and near base strongly recurved, apex acuminate, drying grey green to chestnut brown, darker below, glabrous, closely golden granular glandular below, sparsely granular glandular above, secondary nerves parallel, distinctly curved, especially near margin, c. 1416(20) pairs. Staminate inflorescences sparsely branched 5 cm glabrous to puberulous racemes with 13 cm alternate thread-like divaricate branches some arising from a flower cluster; peduncle to 15 mm; main axis slender, angled; bracts elliptic, 1 mm long, apex acute, puberulous, soon caducous; flower clusters close to touching, in regularly spaced groups of 12; bracteoles minute, ovate, 0.5 mm, apex acute, slightly cucullate, glabrous, erect, persistent, smaller than cluster. Staminate flowers in groups of 4, developing in sequence, shortly pedicellate; stamens 3060, anthers 4-locular. Infructescences delicate glabrous to puberulous, sparsely branched racemes to 7 cm, axes angled, branches thread-like. Fruits regularly, 5 mm distant, single or paired, tiny globose 2 mm, thinly woody, closely golden granular glandular, unarmed or with a few 1 mm slender horns; pedicel slender, 8 mm; calyx tiny, persistent; style papillose, basal, caducous. Seeds globose, verrucose, with sarcotesta.
Distribution Throughout New Guinea (up to Bougainville) and Solomon Islands (New Georgia).
Habitat & Ecology Primary and secondary montane forests, (400)10001800 m.
Note I now believe Corner NGF 13600 (LAE !) from Bougainville belongs here, having found a sheet (Whitmore's Collectors BSIP 3720, K!) from New Georgia.
Macaranga pleytei L.M.Perry, J. Arnold Arb. 34 (1953) 203; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)154; Gen. Macaranga Prodr. (2008) 210. Type: Pleyte 1117 (holo A; iso BO, K, L, SING), Indonesia, Papua, Vogelkop, Sorong, Misool Island near Fakal.
Small trees, 6 m. Twigs slender, 4 mm diam., terete, with long spreading golden hairs, becoming glabrous, blackish brown with conspicuous stipule scars. Stipules united, apparently single, forming a gibbous 4 by 1 cm terminal bud, purplish brown, tomentose with long golden hairs, papery, erect. Leaves: petioles 36 cm, slender, terete, hairy like twigs; blades rotund, c. 12 by 10 cm, leathery, base broadly rounded, 1 cm peltate, lamina with several glands between petiole insertion and margin, margin recurved, apex obtuse or apiculate, drying dark slightly purplish brown above, dull rusty brown below, coarsely bullate between secondary nerves and finely bullate between scalariform tertiaries and quaternaries, above glabrous except for sparse long white hairs, below densely pubescent with spreading white solitary and tufted hairs on all nerves and sparsely black granular glandular, with 3 major and several minor nerves at petiole insertion. Staminate inflorescences slender 14 by 5 cm panicles with 3 axis orders; main axis slender, terete, 1 mm diam.; peduncle 8 cm, sparsely tomentose; branches divaricate, to 2.5 cm, lowest opposite, several with axillary branches, thread-like, slightly zigzag, between tiny, 2 mm evenly spaced flower clusters; bracteoles 2 by 1.5 mm, nerved, slightly fleshy, glabrous, strongly concave, wholly enclosing flower cluster with a slightly peltate obtuse to broadly acute deltoid to subsagittate 1 by 1 mm apex bearing multiple small patellar glands, slightly protruding beyond cluster. Staminate flowers c. 20 per cluster, sessile, minute; stamens 23, anthers 3-locular. Pistillate unknown.
Distribution West New Guinea (known from type only).
Habitat & Ecology Common [in] thinned forest'.
Note Leaves highly distinctive. Staminate bracteoles remarkable.
Macaranga polyadenia Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv add. vi (1919) 25; L.M.Perry, J. Arnold Arb. 34 (1953) 249; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)154; Gen. Macaranga Prodr. (2008) 213, Fig. 17. Syntypes: Ledermann 8664 (K, SING),, Ledermann 8677 (K, SING), both Papua New Guinea, Sepik Region, April River.
Macaranga fimbriata S.Moore, J. Bot. Suppl. 61 (1923) 48; L.M.Perry, J. Arnold Arb. 34 (1953) 248; Airy Shaw, Kew Bull. 31 (1976) 395.
Macaranga fimbriata S.Moore var. doctersii L.M.Perry, J. Arnold Arb. 34 (1953) 249.
Macaranga ovatifolia auct.non Merr.: L.M.Perry, J. Arnold Arb. 34 (1953) 247.
Small trees or to 18m tall and 30 cm diam., sometimes with stilt roots or plank buttresses. Twigs slender, 23 mm diam., at first weakly to strongly angular and usually blackish, at first slightly rusty furfuraceous. Stipules elliptic to ovate-elliptic, c. 10 by 3 mm, apex acuminate, blackish, papery, usually rusty furfuraceous to pubescent, erect, usually mostly soon caducous. Leaves: petioles 611 cm, slender, 1.5 mm diam., terete, blackish, usually glabrous, occasionally rusty furfuraceous, weakly kneed at both ends; blades narrowly to broadly ovate, 915(22) by 410(12) cm, papery, base cuneate to rounded to broadly truncate, sometimes 2 mm peltate, with 2 small glands near petiole insertion, margin (weakly) repand, apex tapering, sometimes abruptly acuminate for 1 cm, drying khaki or brown below dark brown above, glabrous below with close (tiny) granular glands. Staminate inflorescences mostly behind leaves, crowded, patent intermingled dense panicles to 12 cm long, 3(4) axis orders, grey to brown furfuraceous; branches divaricate, secondaries to 6 cm long, tertiaries to 1 cm; peduncle 2 cm; main axis slender, angled and striate; branches opposite or alternate, some sometimes with axillary branches; bracts triangular, to 2 by 1 mm, apex acuminate, usually soon caducous; ultimate branches thread-like; flower clusters grouped at branch tips; bracteoles ovate, 2 by 2 mm, apex acute, persistent, deeply distantly regularly dentate, the teeth sometimes acuminate (subfimbriate), fleshy, both sides densely finely pubescent, erect, longer than cluster. Staminate flowers 5 per cluster, elongate, 1 by 0.5 mm, pedicellate; stamens exserted, 4, anthers 4-locular. Infructescences rather robust, as staminate but only 68 cm long, without axillary branches; secondaries 1 cm distant, to 3 cm long; tertiaries robust, 5 mm. Fruits usually paired, small, 23 mm diam., globose or bilobed (Queensland), very thinly woody, closely black granular glandular; pedicel absent or 1 mm; calyx persistent; stigma bifid, 1 mm, apical, very soon caducous. Seeds globose, very finely wrinkled, with sarcotesta.
Distribution Moluccas (Aru Islands), throughout New Guinea, including Bismarck Archipelago, Admiralty Islands (Manus), Solomon Islands, Santa Cruz Group, Vanuata, Australia (Queensland).
Habitat & Ecology Primary or secondary forest, sometimes riverine or in wet places. Mostly low elevations, rarely to 300600 m. Common.
Notes 1. A polymorphic species but with no discrete variants (Whitmore 1980). The leaf base varies, the markedly truncate state is common in the Bismarck Archipelago, where leaves to 30 cm long also occur. Big leaves are also found on mainland New Guinea. The staminate bracteoles are dentate but subfimbriate when the teeth have elongate tips.
2. Bears a superficial resemblance to M. faiketo. Faiketo is the Kwara'ae name for both species.
3. This differs from other members of the Angustifolia group in its dentate to subfimbriate bracteoles and apical style. I no longer maintain a separate group for it, based on these two characters.
Macaranga praestans Airy Shaw, Kew Bull. 19 (1965) 318; Hook. Ic. Pl. 38 (1974) t. 3718; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 155; Gen. Macaranga Prodr. (2008) 215, Fig. 18. Type: Haviland & Hose 3686K (holo K), Borneo, Sarawak, near Kuching.
Small trees, to 5(10) m. Twigs 3 mm diam., rather sharply angled, glabrous. Stipules subulate, widening at base, 1 cm, glabrous, erect, persistent. Leaves: petioles c. 8 cm long, 2 mm diam., terete, striate, sometimes with spreading 2 mm stiff hairs, kneed at both ends; blades in pseudowhorls of 3 or 4 at 8 cm spacing, elliptic, 1728 by 410 cm, thinly leathery, base cuneate to rounded with a pair of tiny 12 mm convex auricles and 2 conspicuous raised glands, margin undulate often slightly recurved, apically tapering, sometimes abruptly so, to a 24(6) cm caudate tip, drying warm brown, weakly bullate, glabrous with very sparse usually blackish granular glands below, secondary nerves 1215 pairs, sharply raised below, straight, parallel, looping and joining near margin. Staminate inflorescences crowded spikes to 9 cm, arising from a mass of stipule-like bracts, (sparsely) pubescent angular, coarsely striate axis with 5 mm spaced clusters of eventually c. 11 flowers developing in sequence; bracteoles cucullate, ovate, 1 by 1 mm, tapering into a 34 mm caudex. Staminate flowers 1.3 mm diam., developing and falling in sequence, leaving 1 mm pedicel; sepals 3, rounded, glabrous or with a few granular glands; stamens c. 11, filaments 1 mm, anthers 4-locular. Pistillate inflorescences forming 23 cm diam. clusters, spaced along twigs amongst and behind leaves, spikes to 2 cm, axis stout, angled, weakly pubescent and sparsely granular glandular, bearing 16 flowers subtended by 2 stipule-like rudimentary bracts; peduncle 25 mm, granular glandular, with 2 apical leafy pinnately nerved orbicular bracts 2 cm across, blunt or apiculate, living greenish yellow, pink or red, glabrous or with scattered stiff white hairs, enclosing a single flower. Pistillate flower with stout, 3 mm pedicel; sepals ovate, 1 mm, apex acute, glabrous, erect; ovary 1 mm with twin exserted 3 cm smooth twisted filiform caducous styles. Fruits strongly dicoccous, slightly flattened, each lobe 10 by 5 by 4 mm, smooth, densely granular glandular. Seeds not seen.
Distribution Borneo (Sarawak, Brunei).
Habitat & Ecology Primary forest, sometimes peat swamps; locally common and at Lambir in clumps (S.J.Davies, pers. comm.)
Notes 1. Highly distinctive in leaf (especially the base), staminate, and particularly pistillate inflorescence.
2. Dransfield et al. JD 7305, staminate, differs from the type in having the bracteoles oblong, 1.5 by 1.5 mm, with a triangular apex abruptly tipped by a shorter 1 mm caudex. The inflorescences are old, many flowers have fallen leaving protruding pedicels. Despite this difference, on the evidence available, and on JD 7306 pistillate, collected presumably at the same place, this is the same taxon.
Macaranga pruinosa (Miq.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 992; Pax & K. Hoffm. inEngl., Pflanzenr. IV.147.vii (1914) 309; Airy Shaw, Kew Bull. 26 (1972) 289; Whitmore, Tree Fl. Malaya 2 (1973) 106; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 153; S.J.Davies, Harvard Pap. Bot. 6 (2001) 433; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 376, Fig. 17; Whitmore, Gen. Macaranga Prodr. (2008) 217, Fig. 19. Mappa pruinosa Miq., Fl. Ind. Bat. Suppl. Sumatra (1859) 457. Lectotype (designated by S.J.Davies, 2001): Diepenhorst s.n. (U), Indonesia, West Sumatra, Province Priaman.
Macaranga maingayi Hook.f., Fl. Brit. India 5 (1887) 449; Ridl., Fl. Malay. Penins. 3 (1924) 298; Corner, Ways. Trees Malaya (1940) 267. Lectotype (designated by S.J.Davies, 2001): Maingay 1391 (holo: K; iso: L, A [photograph]), Malaysia, Melaka.
Macaranga formicarum Pax & K.Hoffm. in Engl., Pflanzenr. IV147vii (1914) 308; Merrill, J. Straits Br. Roy. As. Soc. (1921) 341 (fomicarium'); Whitmore in Airy SHaw, Kew Bull., Addit. Ser. 4 (1975) 153. Type: Winkler 3025 (not found), Indonesia, Southeast Borneo, Semurung and Sungai Tarik [sic].
Peat swamp tree to 2530 m tall and 3040 cm dbh, flowering smaller; twigs 512 mm in diam., terete, either glabrous and densely glaucous or densely covered in short erect silvery hairs and not glaucous, sometimes hollow and ant-inhabited. Bark pale grey to white, hoop-marked, smooth. Stipules rounded, 815 by 815 mm, erect, usually saccate, glabrous or finely tomentose, the pair not completely encircling the twig, coriaceous, sometimes producing food-bodies on the adaxial surface, usually 35 pairs persistent on the shoots. Leaves: petioles terete, 1025 cm long, sometimes slightly glaucous towards the base, either mostly glabrous with scattered ferrugineus hairs towards the leaf insertion or densely silvery pubescent; blades very broadly ovate to rounded, 825 by 827 cm, evenly trilobed with the central lobe slightly larger than laterals, dissected to c. 1/31/2, deeply peltate to 1.54 cm, base broadly rounded, sometimes slightly cordate, margin ± entire, apices acute or shortly acuminate, adaxial surface mostly glabrous but moderately to densely ferrugineus pubescent along the veins particularly near the petiole insertion, abaxial surface densely pubescent with fine erect silvery hairs and minute ferrugineus hairs, sometimes the silvery hairs absent, minutely furfuraceous; 1° venation palmate with 59 prominent veins, 2° venation scalariform looping and terminating at the margins in prominent conical nectaries, 3° venation densely scalariform, 4° venation weakly scalariform, finest nerves densely reticulate; young leaves adaxial surface densely ferrugineus tomentose, abaxial surface covered in fine erect silvery hairs or a silvery tomentum, the margin often with erect silvery hairs. Staminate inflorescences paniculate, erect, 618 by 410 cm, all axes densely minutely silvery or golden ferrugineus pubescent, the basal axis sometimes ± glabrous, 34 axis orders, main axis short and terete or slightly flattened, first pair of secondary branches alternate without accessory branches; bracts narrowly ovate, 24 by 23 mm, margin entire, apex acute, glabrous or scattered with ferrugineus hairs towards the base and on the margins, persistent; flower clusters with 615 flowers, spirally arranged on the inflorescence branches; bracteoles ovate to narrowly ovate, 35 by 23 mm, often with quite straight sides to almost strap-like, margin shallowly irregularly dentate with 48 small unequal teeth, apex rounded to broadly acute, veins slightly raised on the abaxial surface, both surfaces densely covered with fine silvery and ferrugineus hairs. Staminate flowers c. 0.7 mm long, shortly pedicellate; sepals 3, free, densely ferrugineus pubescent; stamens 23; anthers 4-locular. Pistillate inflorescences paniculate, erect, 518 by 28 cm, sparsely branched, densely ferrugineus pubescent throughout, the hairs slightly less dense on basal axes, 23 axis orders; bracts ovate, 34 mm long, margin entire, glabrous or with scattered hairs on the margins, caducous. Pistillate flowers c. 1.5 mm long; calyx urceolate, densely minutely ferrugineus or golden pubescent; ovary 2-carpellate; styles c. 1 mm long, free, persistent; stigma very shortly dissected. Fruits c. 5 by 56 mm, bilocular, locules subglobose, pedicellate, evenly covered in glandular yellow sticky exudate, without processes; pedicel 46 mm long, finely ferrugineus hairy. Seeds c. 3 mm in diam., lenticular, black, with shallow grooves, encased in a fleshy pale purple aril.
Distribution Peninsular Thailand, Malay Peninsula, Sumatra, Borneo (Central Kalimantan, East Kalimantan, Sarawak). It is abundant in this habitat in Sumatra, the Malay Peninsula and in Borneo, although Davies (2001) has seen no collections of this species from Sabah.
Habitat & Ecology Macaranga pruinosa is a large early successional tree of lowland peat swamp forest. In all three areas it often forms huge monospecific stands in degraded areas. Enormous areas, for example, between KLIA at Sepang and the city of Kuala Lumpur in Peninsula Malaysia, and between Kuching and Kota Samarahan in Sarawak, are completely dominated by this species. The distinctive uniform canopies of M. pruinosa stands can be seen from far away. Little is known about the seedling ecology of this species, but it may prove useful for restoring and limiting erosion in degraded peat swamp areas.
Notes 1. In addition to its very distinctive ecology, M. pruinosa is distinguished from other members of the group by the combination of having deeply peltate leaves, saccate stipules, and relatively short and delicate staminate and pistillate inflorescences.
2. The Malayan and Sumatran plants appear to differ consistently from the Bornean plants as noted by Whitmore (1975). The Malayan/Sumatran specimens often have densely glaucous and glabrous stems, whereas the Bornean specimens usually have finely pubescent and not glaucous young stems. The stems of the Malayan/Sumatran specimens are also often ant-inhabited, whereas those in Borneo are rarely so. Further field work, particularly in the peat swamp forests of southern Borneo, is required to determine whether this variation warrants recognition at the species level.
3. The type specimen of M. formicarum, which Davies (2001) has not been able to find, would also need consideration if the two forms of M. pruinosa were to be recognized as separate taxa. From the description, M. formicarum seems most likely to be M. pruinosa, however the stems are hollow and glaucous unlike much of the Bornean material Davies (2001) has seen.
4. Within Borneo, Davies (2001) suspects that M. pruinosa may hybridize with the closely related M. gigantea. Macaranga gigantea occupies a very wide range of habitats, and it can often be found on the margins of degraded peat swamp forests. Davies (2001) has found several trees in the field and specimens in the herbarium (see for example: Davies 233 from Miri, Sarawak, and Haviland & Hose s.n. from near Kuching) that he suspects may be hybrids. In both cases the leaves and stipules are of intermediate size, and the leaves have shallow dentations. Davies (2001) does not think this is a new and different species, as he has found only a few individuals, and always in areas where M. pruinosa and M. gigantea are locally sympatric.
Macaranga puberula Heine, Feddes Rep. 54 (1951) 232; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 153; S.J.Davies, Harvard Pap. Bot. 6 (2001) 436. Lectotype (designated by S.J.Davies, 2001): Clemens 26526 (holo: A; iso: K, L) Malaysia, Sabah, Dallas.
Submontane tree to 15 m tall and 1520 cm dbh; twigs usually quite stout, 815 mm in diam., greyish, densely pubescent, hairs erect silvery, not glaucous, hollow, housing ants. Bark greyish, smooth, hoop marked. Stipules very broadly ovate to rounded, 1020 by 1025 mm, erect to saccate, finely pubescent at the base to glabrous at the apex, coriaceous, producing food-bodies on the adaxial surface, up to 6 pairs present on the shoots. Leaves: petioles terete, quite stout, 1535 cm long, densely silvery-grey pubescent throughout; blades broadly ovate, 1530 by 1835 cm, deeply trilobed, dissected to 1/22/3 of the leaf length, lateral lobes spreading 1/2 to 4/5 of the size of the central lobe, 13 cm peltate, base broadly rounded, sometimes slightly cordate, margin entire, sometimes with several subprominent conical nectaries along the leaf base, apices shortly acute, adaxial surface glabrous or with scattered erect silvery hairs and minute ferrugineus hairs on the main veins particularly towards the petiole insertion, abaxial surface moderately to densely pubescent with erect silvery hairs and finely furfuraceous; 1° venation palmate with 5 prominent veins and 57 subprominent veins running from the petiole insertion to the basal leaf margin, 2° venation scalariform looping and sometimes terminating at the margins in small conical nectaries, 3°4° venation densely scalariform, finest nerves densely reticulate; young leaves reddish brown, upper surface densely covered in fine ferrugineus hairs, lower surface minutely greyish pubescent. Staminate inflorescences paniculate, erect, 2030 by c. 20 cm, all axes densely covered in erect silvery hairs, to 4 axis orders, main axis flattened, first pair of secondary branches ± opposite usually with accessory branches; bracts narrowly ovate, 48 by 35 mm, margin entire, apex acute, glabrous or with scattered silvery hairs on the margins and towards the base, persistent; flower clusters with c. 1015 flowers, spirally arranged on inflorescence branches; bracteoles ovate-elliptic, 2.55 by 24 mm, enclosing flower clusters, margin irregularly dissected, apex acute to acuminate, veins slightly raised on abaxial surface, both surfaces densely covered with minute silvery to reddish-brown hairs. Staminate flowers c. 1 mm long, pedicellate; sepals free, apex densely covered in minute red-brown hairs; stamens 23; anthers 4-locular. Pistillate inflorescences paniculate, erect, 1535 by 1020 cm, stout, densely pubescent throughout with long silvery-grey hairs, up to 4 axis orders, bracts caducous, not seen. Pistillate flowers c. 1.5 mm long, solitary in bract axils; calyx urceolate, c. 1 mm long, apex scattered with fine silvery hairs, persistent; ovary 2-carpellate, c. 1 mm long; styles c. 1 mm long, free, persistent; stigma not dissected. Fruits 34 by 45 mm, bilocular, locules globose, pedicellate, evenly covered in sticky glandular exudate, without processes; pedicel 47 mm long, densely finely pubescent. Seeds c. 22.5 mm in diam., lenticular, black, with shallow grooves, encased in a fleshy pale purple aril.
Distribution Borneo (Brunei, Sabah, Sarawak). This species is known from Gunung Kinabalu in Sabah in the north, through the mountains of Brunei, to the Kelabit Highlands in the Sarawak in the south. It is probable that this species is more widespread in submontane forest in Kalimantan and Sarawak.
Habitat & Ecology A submontane early successional tree known from 8001800 m. This species occurs in openings in primary forest, and in degraded areas. The tree has a dark grey-green appearance.
Notes 1. The combination of densely pubescent non-glaucous stout twigs, pubescent inflorescence axes, and peltate leaves distinguish M. puberula from its close relatives.
2. This species is most closely related to M. pearsonii and M. pruinosa. In contrast with M. puberula, both M. pruinosa, a species of peat swamp forests, and M. pearsonii, a species of dryland forests, are restricted to the lowlands. Macaranga pearsonii has densely glaucous and glabrous stems. Macaranga pruinosa has much more slender twigs, smaller leaves, and smaller and less well branched staminate and pistillate inflorescences than M. puberula.
Macaranga punctata K.Schum., Fl. Kais. Wilh. Land (1889) 80; K.Schum. & Lauterb., Fl. Schutzgeb. Sόdsee (1900) 397; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 362; L.M.Perry, J. Arnold Arb. 34 (1953) 227; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)154; Gen. Macaranga Prodr. (2008) 217. Type: Hollrung 825 (iso: BO, K, P), Papua New Guinea, Sepik Area, Malu, Augusta Station.
Macaranga isadenia Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 377.
Macaranga ovalifolia Pax & K.Hoffm. in Engl.,Pflanzenr. IV.147.xiv add. vi (1919) 29; non Ridley (= M. papuana J.J.Sm.).
Macaranga pseudopeltata Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv add. vi (1919) 29.
Macaranga maluensis Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv add. vii (1924)185.
Macaranga punctata K.Schum. var. whitei L.M.Perry, J. Arnold Arb. 34 (1953) 229.
var. punctata
var. punctata: Whitmore, Gen. Macaranga Prodr. (2008) 217. Type: as the species.
Shrubs or small trees, 2(12) m, occasionally sarmentose. Twigs slender, 23 mm diam., broadly angled, later terete, finely rusty furfurescent, rarely glabrous, dark brown. Stipules (narrowly) oblong-ovate, 3 by 1.52 mm, apex broadly acute, thin, pubescent, erect, soon caducous, usually leaving a conspicuous scar. Leaves: petioles c. 5 cm, slender, 1.5 mm diam., terete, sparsely to densely pubescent, not kneed; blades ovate to elliptic, 1423 by 513 cm, thinly leathery, base rounded, broadly rounded or rarely broadly cuneate, cordulate or to 5 mm peltate, with several usually small conspicuous glands and 3 main and usually additional finer nerves at petiole insertion, apex acute or usually abruptly 15 mm acuminate, drying khaki, below midrib and secondary nerves usually darker than lamina, above glabrous or rarely pubescent on lower midrib, below blackish granular glandular and usually rusty pubescent (rarely glabrous) on nerves; tertiaries scalariform, conspicuous below. Staminate inflorescences open racemes or panicles to 13 cm long, 2 or 3 axis orders, furfuraceous (occasionally glabrous) branches divaricate, secondaries to 8 cm, 14 cm apart; peduncle 25 cm; main axis slender, 12 mm diam., usually terete, occasionally slightly flattened, branches alternate or rarely opposite, arising from a flower cluster, sometimes with axillary branches; bracts not seen; ultimate branches thread-like to 2 cm; flower clusters 1 cm apart, evenly spaced; bracteoles distally triangular to oblong, 2 by 2 mm, apex acute, persistent, spreading, longer than cluster, thick, fleshy, entire, with 26 sunken patellar glands, plus a variously developed concave base bearing the flowers, hairy outside. Staminate flowers in clusters of 811, maturing sequentially, shortly pedicellate; sepals free, with a few granular glands; stamens 47, long-exserted at maturity, anthers 4-locular. Infructescences short spikes or less often racemes to 7(10) cm, axis flattened, furfuraceous; peduncle 1.56 cm; bracts elliptic, 5 mm, leaf-like, rusty pubescent, soon caducous. Fruits solitary, widespaced along axis, plus a terminal group of 24, round, 6 mm diam., green, drying black, thinly woody, with dense, 3 mm sparsely hairy spines, caducous to leave bald patches, granular glandular; pedicel 1 mm, stout; calyx small, persistent; styles 2 mm, recurved, plumose, caducous. Seeds round, 5 mm, coarsely verrucose, with sarcotesta.
Distribution Throughout New Guinea.
Habitat & Ecology Primary forest, sometimes riverine, lowlands and once at 990 m.
Notes 1. Somewhat polymorphic. Distinctive in the staminate inflorescence branches arising from flower clusters, multinerved gland-bearing leaf base, leaves with dark nerves, fruits spread along the infructescence and with short somewhat caducous spines.
2. van der Sidje BW 4045 from Jayapura is unusual in being entirely glabrous.
var. subulata Whitmore
var. subulata Whitmore, Gen. Macaranga Prodr. (2008) 219. Type: P. van Royen & Sleumer 6190 (holo K; iso G, P), Indonesia, Papua, Cycloop Mountains, road Hollandia to Sentani.
Differs in stipules 7 mm long, subulate.
Distribution West New Guinea (known from type only).
Habitat & Ecology Secondary forest, 70 m alt.
Note Apart from the stipules this fits well into M. punctata. Like a few other collections it is glabrous.
Macaranga puncticulata Gage, Rec. Bot. Surv. India 9 (1922) 246; Ridl., Fl. Malay. Penins. 3 (1924) 302; Whitmore, Gen. Macaranga Prodr. (2008) 219, Fig. 20. Type: Ridley 13259 (holo CAL?; iso K, SING), Peninsular Malaysia, Johore, Tempayan River.
Macaranga caladiifolia auct. non Gage: Whitmore, Malay. Nat. J. 20 (1967) 98; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 150.
var. puncticulata
var. puncticulata: Whitmore, Gen. Macaranga Prodr. (2008) 219, Fig. 20. Type: as the species.
Small trees, up to 10 m tall. Twigs usually hollow, ant-inhabited, stout, to 1 cm diam., or sometimes solid, slender, 3 mm diam., terete, smooth or coarsely striate, with conspicuous circular stipule scars, usually glabrous, sometimes glaucous, sometimes furfuraceous. Stipules elliptic, 7 by 2 mm, apex acuminate, dark brown, papery, glabrous, furfuraceous or weakly tomentose, erect, soon caducous. Leaves: petioles 612 cm, slender, terete, striate, glabrous, glaucous, with a furfuraceous knee at top; blades ovate-oblong, 1218 by 812 cm, thinly coriaceous, base broadly rounded, 24 cm peltate, margin with distant protruding small glandular teeth, often slightly sinuate, sometimes recurved, apex 1 cm acuminate, drying khaki brown, below very densely puncticulate and glaucous, midrib and main nerves furfuraceous to glabrous. Staminate inflorescences small sparsely branched panicles, to 8 cm, finely rusty pubescent, 3 orders of axis; main branches alternate, divaricate; peduncle to 5 cm, slightly flattened; bracts ovate-oblong, 4 by 2 mm, caducous; flower clusters crowded on 5 mm tertiary axes; bracteoles overlapping, erect, persistent, ovate, 3 mm, concave, tapering into a 3 mm acumen, rusty furfuraceous, entire. Staminate flowers in clusters of c. 6, 0.5 mm with 0.5 mm pedicel; sepals hairy; stamens 2; anthers 4-locular. Infructescences spicate, 38 cm, peduncle 34 cm. Fruits a few, crowded at apex, globose to depressed globose, to 15 mm diam., black, leathery, closely granular glandular; pedicel stout, 3 mm, rusty furfuraceous; sepals persistent, 2 mm, rusty furfuraceous; stigmas apical, small, 1 mm. Seeds not seen.
Distribution Malay Peninsula, Borneo (Sarawak; West Kalimantan: Palung).
Habitat & Ecology Lowlands, peat swamp and permanent and periodic freshwater swamp forest, occasionally dry land. Sometimes ant inhabited.
Notes 1. The Kalimantan collection, Sidiyasa 1428 (K!), has the fruiting pedicel 10 mm.
2. The leaves have the under surface puncticulate rather than granular glandular. Similar to M. pachyphylla var. glabra, but is distinct in its lowland and nearly always swampy habitat, elliptic stipules, sparsely branched staminate inflorescences and shorter very stout fruiting pedicel.
var. tenuiramea (Pax & K.Hoffm.) Whitmore
var. tenuiramea (Pax & K.Hoffm.) Whitmore, Gen. Macaranga Prodr. (2008) 220. Macaranga tenuiramea Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 384; Merr., J. Str. Br. Roy. Soc. Spec. No. (1921) 342. Type: Haviland 464 (iso: CGE, K, P, SAR, W), Borneo, Sarawak.
Macaranga caladiifolia auct. non Becc.: Whitmore, Tree Fl. Malaya 2 (1973) 111; in Airy Shaw, Kew Bull. 36 (1981) 319.
Differs in Twigs always hollow, unevenly grossly swollen; Leaves: blades narrowly oblong, with truncate base, 813 by 36 cm, usually (strongly) glaucous below, margin with very conspicuous protruding distant glandular teeth.
Distribution Malay Peninsula (once collected), Sumatra, Borneo (Sarawak).
Habitat & Ecology Lowlands. Uncommon.
Notes 1. An extreme form of M. puncticulata, but strikingly different in leaf shape from var. puncticulata to which it is linked by a few intermediate collections. Ngadiman SFN 36636 approaches this variety. A Bornean endemic except for one collection (Whitmore KEP FRI 3825 from Pahang).
2. Easily confused, especially in the herbarium, with M. caladiifolia (in S.J.Davies 2001: 393) in its two distinctive features, namely its leaf shape and twig. However, M. caladiifolia has distinctive and bizarre secondary nervation, is completely glabrous, not furfuraceous, and has different inflorescence, flower and fruit characters.
3. J.A.R.Anderson and I confused this with M. caladiifolia (published 1914) to which in consequence I (Whitmore 1975) reduced M. puncticulata (1922).
Macaranga quadriglandulosa Warb., Bot. Jarb. 13 (1891) 350; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 356; J.J.Sm., Nova Guinea Bot. 12 (1917) 546; Mansfeld, J. Arnold Arb.10 (1929) 232; Kaneh. & Hatus., Bot. Mag. Tokyo 52 (1938) 411; L.M.Perry, J. Arnold Arb. 34 (1953) 237; Airy Shaw, Kew Bull. 23 (1969) 98; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 155; Gen. Macaranga Prodr. (2008) 222. Macaranga quadriglandulosa Warb.var. genuina Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii(1914) 356, nom. illeg. Type: Warburg 20534 (iso A), apua New Guinea, New Britain, Kerawara and Mioko Islands.
Macaranga tanarius (L.) Mόll.Arg. var. abbreviata J.J.Sm., Nova Guinea Bot. 8 (1912) 238, 791. Macaranga quadriglandulosa Warb. var. abbreviata (J.J.Sm.) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 356.
Macaranga quadriglandulosa Warb. var. digyna Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 356.
var. quadriglandulosa
var. quadriglandulosa: Whitmore, Gen. Macaranga Prodr. (2008) 223. Type: as the species.
Small trees to 10 m. Twigs 5 mm diam., very strongly angled, glabrous or at first sparsely pilose with long white hairs, occasionally slightly glaucous. Stipules elliptic, to 15 mm long, 4 mm wide at base, tapering to a long tip, dull brown, papery, sparsely pilose or glabrous on both surfaces, spreading to recurved, subpersistent. Leaves: petioles (6)18 cm, slender (1)1.5(2) mm diam., terete, sparsely pilose or glabrous, narrow knee at base, knee at top; blades round to ovate-oblong, size very variable, (1014(23) by (6)12(18) cm, drying papery and brittle and usually crinkled, base strongly (14 cm) peltate, broadly rounded, lower main nerves usually with large flat glands totalling 38, but sometimes only 1 or none, margin regularly finely glandular dentate, apex acuminate, drying pale brown, sometimes greenish above, below main nerves sparsely pilose, with spider's web nervation. Staminate inflorescences dense panicles to 24 by 10 cm, 3 axis orders, pubescent to glabrous, branches ascending; peduncle 47 cm; main axis stout, flattened, to 3 mm across; lower branches usually opposite, usually with axillary branches; bracts not seen; flower clusters evenly c. 3 mm spaced; bracteoles persistent, larger than flower clusters, very strongly concave, 3 by 4 mm, apex roundish, papery, reticulately nerved, outside granular glandular, margin entire with a few glands. Staminate flowers 7 per cluster, sessile; stamens 2, anthers 4-locular. Infructescences (2)9 cm spikes tipped by a single or rarely 45(8) fruit(s), axis variously flattened, to 3 mm diam., furfuraceous to glabrous. Fruits usually very weakly bilobed, to 12 by 18 mm, or globular or to 5 locular, closely beset with fine sinuous spines, body and sometimes spines yellowish granular glandular; pedicel stout, 8 mm glabrous; calyx persistent. Seeds round, coarsely rugose, with sarcotesta.
Distribution Throughout New Guinea, Bismarck Archipelago, Solomon Islands.
Habitat & Ecology Secondary forests lowlands to 900 m. Common.
Note The eponymous leaf glands not always present or easy to see.
var. variabilis L.M.Perry
var. variabilis L.M.Perry, J. Arnold Arb. 34 (1953) 239; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)155; Gen. Macaranga Prodr. (2008) 222. Type: Brass 1204 (holo A; iso K, P), Papua New Guinea, Central Prov., Kerema.
Differs solely in the leaves not peltate.
Distribution East New Guinea.
Macaranga racemohispida Whitmore, Gen. Macaranga Prodr. (2008) 224. Type: De Vogel 9785 (holo L; iso BO, K, MAN), Indonesia, Papua, Sorong, Ayawasi Valley.
Similar to M. hispida but easily told apart from the following characters: Twigs slightly glaucous, almost glabrous with just a few sparse hairs. Leaves: petioles 27 cm, flattened above near base, distantly shallowly striate, glabrous except for a few hairs, weakly glaucous; blades very big, 40 by 34 cm with the basal sinus very deep, 11 cm, lower surface granular glands golden. Staminate unknown. Infructescences 19 cm narrow racemes, main axis ornament as petioles; peduncle 7 cm; branches to 5 cm, ascending, tomentose. Fruits single or paired, irregularly spaced and at branch tips, crowded, 23 lobed, 8 by 10 mm (? immature) with dense soft hairy caducous 3 mm processes, closely golden granular glandular; pedicel to 14 mm, tomentose; calyx persistent.
Distribution West New Guinea (known from type only).
Habitat & Ecology Secondary forest on deep coarse white sand, 450 m.
Uses Leaves used with sago and kino to seal letters.
Macaranga rarispina Whitmore, Kew Bull. 29 (1974) 450; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 155; Gen. Macaranga Prodr. (2008) 226. Type: Daud & Tachun SFN 35721 (holo K; iso L), Borneo, Sarawak, Sungai Temulan.
Aff. M. anceps Airy Shaw': Airy Shaw, Kew Bull. 19 (1965) 324.
Trees, 4(13) m. Twigs 3 mm, angled, smooth ageing rugose, warm brown, arachnoid pubescent or glabrous. Stipules and bud scales subulate, 10 mm, dull brown, erect, glabrous, crowded at tip, subpersistent at nodes near twig tip. Leaves: petioles 612 cm, slender, 2 mm diam., terete, coarsely striate, glabrous, with a conspicuous apical knee; blades elliptic 1625 by 611 cm, thinly leathery, base rounded or cuneate, with conspicuous gland-bearing auricles 15 mm long, margin usually sinuous and with distant small glandular teeth, apex fairly abruptly 1015 mm acuminate, drying khaki fawn, glabrous with neither puncticulations nor granular glands. Staminate inflorescences apical, crowded 5 cm catkin-like spikes covered in a diffuse arachnoid pubescence; axis not visible; bracteoles ovate, 2 by 2 mm, apex rounded, spreading, papery, margin entire, persistent, outside arachnoid pubescent, shorter than the closely touching clusters of 4 flowers. Staminate flowers developing in sequence; stamens c. 10, anthers 4-locular. Infructescences crowded, 58 cm spikes; axis stout, 2 mm diam., more or less terete, striate, pubescent or glabrous, fruits crowded in apical 1 cm; bracts not seen; bracteoles subulate, 4 mm, with a small 1 mm ovate base, sparsely pubescent, erect, caducous. Fruits strongly bilobed, each lobe 10 by 10 mm, woody, densely tawny pubescent with sparse 12 mm stout spines; pedicel stout, 11 mm, rugose, pubescent; sepals elliptic, 5 mm, pubescent; styles not seen. Seeds 9 by 10 mm, smooth, with a thin sarcotesta.
Distribution Throughout Borneo.
Habitat & Ecology Primary lowland forest, once on ultrabasics.
Note This has a superficial resemblance to M. anceps but differs in many respects, notably stouter petioles, larger leaves, longer stipules and buds, and larger fruits on longer pedicels.
Macaranga recurvata Gage, Rec. Bot. Surv. India 9 (1922) 246; Ridl., Fl. Malay. Penins. 3 (1924) 301; Whitmore,Tree Fl. Malaya 2 (1973) 111, t.8; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 149; Gen. Macaranga Prodr. (2008) 226. Syntypes: Ridley s.n. (SING), Singapore, Bukit Timah; Ridley 10420 (SING), Singapore, Bukit Manday; Ridley 10934 (K, SING), Singapore, Mandai.
Trees to 30 m tall, 50 cm diam. Twigs solid, near tip drying angled, 4 mm diam., terete behind the tip and ultimately very stout, 15 mm diam., with a well developed pale striate bark, glabrous, tip dark brown. Stipules ovate-oblong, to 25(40) by 10(15) mm, apex acute, narrowing slightly at base, often cucullate, blackish brown, papery, glabrous, erect, fairly persistent. Leaves: petioles 1620 cm, stout, 3 mm diam. round to angled, striate, glabrous, weakly kneed at both ends; blades ovate-oblong, 2240(45) by 1626(34) cm, leathery, base broadly rounded, c. 5 cm peltate, with spider's web nervation, margin slightly recurved, weakly crenate and distantly glandular dentate, apex acute, usually drying chestnut brown, below slightly glaucous, glabrous and finely closely golden granular glandular, secondary nerves straight, darker than lamina, tertiaries faint, distant, scalariform. Staminate inflorescences fairly dense panicles 10(20) cm long, 3 axis orders, glabrous; secondary branches divaricate; peduncle 510(20) cm, 45 mm wide; main axis stout, flattened, 24 mm wide, striate; lower branches opposite, secondaries to 6 cm, sinuous; bracts oblong, to 5 by 3 mm, apex acute, papery, spreading, somewhat cucullate, subpersistent; flower clusters tipping tertiary and some secondary branches, as catkins, 12 cm long, with a 1 cm stalk; bracteoles erect, overlapping, to 6 by 3 mm, broadest at 3.5 mm, or smaller, tapering to acute apex, sometimes coarsely toothed on upper part, leathery, glabrous, folded with a 2 by 2 mm basal (? glandular) patch. Staminate flowers in clusters of 57, 1 mm diam., pedicel 1 mm; stamens 23, anthers 4-locular. Infructescences similar to staminate inflorescences but sparsely branched 7 cm racemes with a few distant, 1 cm short secondary branches. Fruits solitary at 1 cm spacing, usually dicoccous, 5 by 10 mm, or tricoccous, 7 by 11 mm, cocci splitting, thinly woody, black granular glandular; pedicel stout, 3 mm, glabrous; sepals thick, 1 mm, persistent; styles 2, 1 mm, apical recurved. Seeds round, 4 mm, coarsely pitted, with sarcotesta.
Distribution Malay Peninsula, Sumatra, East Java, Borneo, Lesser Sunda Islands (Bali).
Habitat & Ecology Primary (Agathis) and secondary lowland forest up to 500 m (or at 1700 m in Bali). Often swampy places. On acid, sandy soil.
Note In essence a more robust version of M. diepenhorstii from which it differs mainly in usually very stout twigs, larger stipules, leaves larger, leathery, with straight dark secondary nerves, staminate inflorescence bracts subpersistent, longer catkins, bracteole shape, and shorter stout fruiting pedicel.
Macaranga reiteriana Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xvii add. vii (1924) 185; L.M.Perry, J. Arnold Arb. 34 (1953) 201; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 156; Gen. Macaranga Prodr. (2008) 227. Syntypes: Ledermann 11811, 11979, 12063a, 12159 (not seen), Papua New Guinea, Madang Prov., Schraderberg.
Small trees, 8 m. Twigs slender, terete, 23 mm diam., at first tawny tomentose, soon glabrous, with conspicuous stipule scars. Stipules united, apparently single, forming a sinuous, elliptic, 7 by 0.3 cm terminal bud. Leaves: petioles 2 cm, slender, 1 mm diam., tawny tomentose, kneed at both ends; blades elliptic, 813 by 22.8 cm, papery, base rounded with 12 inconspicuous glands, margin revolute, narrowing fairly abruptly into 15 mm caudex, drying grey above, below pale brown with close to sparse black granular glands, glabrous above, below tomentose to glabrous on midrib. Staminate inflorescences delicate, 8 cm racemes; axis slender, 1 mm diam., terete, sericeous; peduncle 4 cm; branches thread-like to 1 cm, zigzag between evenly spaced distant tiny flower clusters; bracteoles spreading, roundish, 1 by 2 mm, glabrous, bearing a 30 flowered cluster and with an acute hastate, 1.5 by 1.2 mm erect terminal limb, fleshy, bearing many adaxial glands, margin crenate, larger than cluster. Staminate flowers minute, subsessile; stamens 3, anthers 3-locular. Infructescences delicate 8 cm spikes with 0.8 mm diam. terete tomentose axis and a single globose terminal fruit. Fruits 5 mm diam. (? immature), spineless, granular glandular, sessile; sepals caducous, 2 by 2 mm, apex acute. Seeds unknown.
Distribution Throughout New Guinea
Habitat & Ecology Secondary forest, 14002300 m.
Notes 1. Vinas & Wiakabu LAE 59525 differs in being wholly glabrous but otherwise matches.
2. Named for Kathe Reiter, expert on the flora of Silesia.
Macaranga rhizinoides (Blume) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1011; J.J.Sm., Meded. Dep. Landbouw. 10 (Bijdr. 12) (1910) 485; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 348; Backer & Bakh.f., Fl. Java 1 (1963) 487; Whitmore, Gen. Macaranga Prodr. (2008) 228. Zanthoxylum rhizinoides Blume, Bijdr. (1825) 248; Miq., Fl. Ind. Bat. (1859) 669. Syntypes: Blume s.n. (L), Java, montis Burangrang; Unknown s.n. (L), Java.
Mappa denticulata Blume var. tomentosa Moritzi ex Zoll., Linnaea 28 (1856) 308; Verh. Natuurk. Ver. Ned.-Indiλ 1(1856) 9.
[Macaranga blumeana Mόll.Arg. ex Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 349, nomen nudum.]
Trees 1540 m; young parts densely crisped rufous pubescent. Twigs 3 mm diam., angular, sometimes strongly coarsely ridged. Stipules ovate, 10 by 3 mm, apex acute, densely rufous pubescent, leathery, becoming recurved, caducous. Leaves: petioles 912 cm, slender, 2 mm diam., weakly crisped pubescent, constricted near base; blades broadly triangular-ovate to ovate 1220 by 87 cm, leathery, base broadly rounded to truncate, to 11.5 cm peltate or sometimes almost cordate, margin coarsely shallowly irregularly crenate, below distantly glandular denticulate, apex acute, with an evanescent dense ferruginous tomentum, under this all nerves (and more densely the lamina) with a white pubescence, hairs sometimes tufted, lamina black granular glandular, closely strongly reticulate, above becoming glabrous. Staminate inflorescences lax pyramidal 5 11 cm racemes; peduncle 2.5 cm; branches flexuous, spreading, rusty tomentose; bracts 27 mm, contracted at base, lightly dentate, tomentose, caducous; bracteoles fairly narrow triangular-ovate, 23 mm long, entire or denticulate, more or less concave. Staminate flowers in dense clusters of 10, c. 1.5 mm diam.; sepals 2 or 3, tomentose outside; stamens 714, anthers (3)4-locular. Pistillate inflorescences racemes, 9 by 6 cm; peduncle somewhat flattened, 3 mm diam., densely rusty pubescent; branches normal to axis, to 3 cm, slightly zigzag, sometimes arising from a flower cluster; bracts a few, persistent, triangular, 2 mm long, apex acute, entire, rufous pubescent, recurved. Pistillate flowers: calyx urceolate, shortly lobed, enclosing ovary; latter 1-locular, sparsely viscid granular glandular, glabrous. Fruits sometimes paired, mostly single, obliquely globose, 3 mm, smooth, thinly woody; pedicel 5 mm; sepals 2, triangular, 1 mm long, persistent; stigma small, plate-like, apical, caducous. Seeds round, smooth, rugulose, with sarcotesta.
Distribution Sumatra (uncommon), Java (abundant).
Notes 1. Abundant at BO but poorly represented at K and L.
2. The description is partly paraphrased from Pax & Hoffmann (1914).
Macaranga rorokae Whitmore, Gen. Macaranga Prodr. (2008) 229. Type: Van Balgooy 3351 (holo K; iso BO, L), Sulawesi, Sopu Valley, Mt. Roroka Timbu.
Small trees, 20 m. Twigs angular, 5 mm diam., coarsely striate, finely rufous tomentose. Stipules ovate-elliptic, to 12 by 4 mm, apex acuminate, thinly leathery, tomentum as twigs, erect, soon caducous. Leaves: petiole 18 cm, round, finely rufous tomentose; blades ovate, 1725 by 1523 cm, leathery, base broadly truncate, sometimes very broadly shallowly cordate or 5(15) mm peltate, with 2 or more glands near petiole insertion, margin with distant protruding glands, apex acute, drying rufous brown below, very weakly bullate in parts, below the main nerves and reticulations variously raised, velvety with dense rufous pubescence and dark conspicuous or inconspicuous granular glands. Staminate inflorescences dense narrow velvety racemes 8 cm long; branches alternate, distant to 4 cm, lower ones divaricate; peduncle 1 cm; main axis stout, 1.5 mm diam., terete, mid inflorescence with a few axillary branches; main axis bracts elliptic, 3 by 1 mm, thick, velvety mostly caducous; branches slender; flower clusters crowded, globose, 3 mm across, subtended by persistent bracteoles; latters erect, deltoid, 2 by 3 mm, apex acute, entire, velvety outside, glabrous inside, shorter than clusters; these with c. 18 flowers on short axis in 23 subgroups subtended by deltoid bracteoles, 1.5 by 1 mm or less, hairy outside, glabrous inside. Staminate flowers 1 mm diam., sessile; sepals 3, hairy outside, glabrous inside; stamens 8, anthers 4-locular. Pistillate inflorescences narrow sparsely branched racemes to 7 cm, branches 1.5 cm, axes rufous pubescent; bracts and bracteoles persistent, ovate, 2 by 1 mm, apex acute or obtuse, very thick, pubescent outside, glabrous inside. Young Fruits in clusters of 3, closely granular glandular; pedicel long, 4 mm, rufous; calyx persistent; styles 2, erect, 5 mm, brittle, therefore caducous. Seeds unknown.
Distribution Sulawesi.
Notes 1. First record of a member of the Brunneo-floccosa' group outside New Guinea. The twin apical styles are distinctive, as are the (slightly) bullate pubescent leaves with dark granular glands, short racemes and long-pedicellate fruits. Closest to M. trichanthera but clearly different. The structure of the staminate flower clusters is unique.
2. The epithet is derived from the type locality.
3. Teysmann HB 12491 has the leaves 1015 mm peltate and the reticulations only weakly raised below. It is sterile and conceivably a juvenile.
Macaranga rostrata Heine, Feddes Rep. 54 (1951) 233; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 152; S.J.Davies, Harvard Pap. Bot. 6 (2001) 416. Lectotype (designated by S.J.Davies, 2001): Clemens 34459 (holo: A; iso: K, L), Malaysia, Sabah, Colombon Basin, Gunung Kinabalu.
Submontane tree to 20 m tall; twigs slender, 35 mm in diam., terete, glabrous or finely pubescent with short silvery hairs, not glaucous, solid, not housing ants. Stipules ovate-elliptic, 510 by 35 mm, erect and spreading from the twig and sometimes slightly recurved when dry, abaxial surface finely pubescent along the midrib and towards the base, adaxial surface mostly glabrous, the pair not completely encircling the twig, leathery but not succulent, 14 pairs present on shoots. Leaves: petioles terete, 312 cm long, finely pubescent with short silvery or golden hairs, rarely almost glabrous; blades ovate-elliptic, broadest near the base, 920 by 410(12) cm, not or only shallowly peltate with the petiole insertion <15 mm from the basal leaf margin, base rounded or slightly cordate when not peltate, margin entire to almost wavy or serrate, apix acute to narrowly acuminate, adaxial surface glabrous or rarely with minute scattered hairs along the veins and with 24 round or oval c. 1 mm diameter flat nectaries adjacent to the basal leaf margin near the petiole insertion, abaxial surface densely pubescent along all veins or rarely sparsely pubescent, not glaucous; 1° venation palmate with 57 prominent veins, 2° veins scalariform looping near the leaf margins and terminating in small protruding flattened nectaries, 3° veins scalariform; young leaves red-brown, densely pubescent along the veins, the very youngest leaf bud densely covered in erect pointed silvery or golden hairs. Staminate inflorescences slender, green when fresh, drying blackish, paniculate, erect, 415 by 16 cm, glabrous or sometimes scattered with silvery hairs on the basal axes and minute ferrugineus hairs on the distal axes, to 3(4) axis orders, main basal axis short, flattened, first pair of secondary branches ± opposite and usually with accessory branches; bracts elliptic, 512 by 36 mm, margin entire, apex acute, mostly glabrous or margin and adaxial surface scattered with minute ferrugineus hairs, caducous; flower clusters with 1525 flowers, spirally arranged and evenly spaced; bracteoles broadly rounded, 24 by 35 mm, cupulate, enclosing flower clusters, margin entire, apex shortly acute, adaxial surface densely covered with minute ferrugineus hairs particularly towards the base, abaxial surface scattered with ferrugineus hairs and yellow granular glands. Staminate flowers c. 1 mm long, sessile; sepals fused, splitting to c. 1/5, apex densely covered in minute red-brown hairs; stamens 1; anthers 3-locular. Pistillate inflorescences erect, shortly paniculate, 37 by 13 cm, stout, 12 axis orders, secondary branches ± opposite, green when fresh and drying black, mostly glabrous but main basal axis usually with scattered erect silvery hairs; bracts caducous, not seen. Pistillate flowers 56 mm long, solitary; calyx c. 3 mm long, urceolate, glabrous, persistent, splitting irregularly as ovary expands; ovary 4(5)-carpellate, 34 mm long; styles 23 mm long, fused at base, free and spreading from c. 1/3 of length, persisting to form a prominent 34 mm long crown at fruit apex; stigma not dissected. Fruits subglobose, 68 (excluding horns) by 910 mm, sessile, one discrete glandular patch on each carpel wall developing into a very long and slender horn-like process 715 mm long, covered in sticky yellow-green exudate. Seeds subtriangular-ovoid, c. 4 mm in diam., black, with shallow coarse grooves and a small cruciform scar, encased in a fleshy aril, aril color not seen.
Distribution Borneo (Sabah and Sarawak). This species is endemic to north Borneo; distributed from Gunung Kinabalu in the north, through the Crocker Range in Sabah, to the Kelabit Highlands in Sarawak. It seems likely that it also occurs further south in Sarawak and in the neighboring mountains of Kalimantan, although it is yet to be collected there.
Habitat & Ecology Present in submontane forests between 800 and 2300 m. It occurs both in undisturbed forest and on forest margins.
Notes 1. Macaranga rostrata is easily recognized by having entire leaves, spreading stipules, solid twigs that are not inhabited by ants, and exceptionally long horns on the mature fruits. In addition, the relatively high altitude inhabited by this species is unusual for section Pachystemon.
2. Despite these differences M. rostrata shares unistaminate flowers and 45-carpellate fruits with the other species of the largest clade of section Pachystemon. It is probably most closely related to M. depressa f. strigosa, with which it shares relatively small spreading stipules, staminate bracteoles with a shortly acute to acuminate apex, solid stems, and exceptionally long horns on the fruits. It differs from that species in having evenly pubescent to glabrous twigs and young shoots, and in having entire leaves; M. depressa f. strigosa has densely strigose hairs on the shoots and shallowly to deeply trilobed leaves.
Macaranga rufescens S.J.Davies, Harvard Pap. Bot. 6 (2001) 437. Type: Davies 229 (holotype: A), Malaysia, Sarawak, Sen Sen logging concession, between Miri and Lambir Hills National Park, 4°20'N 113°50'E.
Large tree with a reddish-green appearance, to 2030(35) m tall and 3040(50) cm dbh, although flowering from 35 m in open areas; twigs 815 mm in diam., terete, glabrous, sparsely to moderately glaucous. Bark pale grey, white or light brown, prominently hoop-marked, smooth or minutely fissured. Stipules large, orbicular and flap-like, 1220 by 1220 mm, surrounding the stem, erect from the stem, numerous pairs persistent well down the stem, coriaceous, glabrous, producing food-bodies on the adaxial surface. Leaves: petioles terete, slender, 1040 cm long, slightly glaucous towards the base, glabrous, usually red when fresh; blades broader than long to almost rounded, not flat, slightly convex, 1425 by 1435 cm, deeply trilobed, dissected to c. 1/23/4, the lateral lobes almost the same size as the central lobe, the lateral lobes slightly asymmetrical, not peltate or rarely 15 mm peltate, base very broadly rounded, strongly cordate, margin ± entire with small conical nectaries scattered along the distal margin but almost always absent from the basal margin, apices acute, adaxial surface glabrous or with scattered ferrugineus hairs near the petiole insertion, abaxial surface densely grey-brown furfuraceous and rarely with scattered erect silvery hairs along veins, 1° venation palmate with 5 prominent veins and 23 less prominent veins arising from the petiole insertion, 2°3° veins densely scalariform, 4° venation scalariform, minor veins usually prominent and finely reticulate; young leaves reddish, adaxial surface densely covered in minute ferrugineus hairs, abaxial surface densely furfuraceous and rarely with scattered erect sharp hairs. Staminate inflorescences erect narrow panicles usually not extending above the leaves, 1025 cm long, to 4 axis orders, basal unbranched axis slightly flattened and less than 1/2 of the total inflorescence length, first pair of secondary branches alternate without accessory branches, basal axes glabrous, distal axes ferrugineus pubescent; bracts ovate, 515 mm long, margin entire, glabrous or rarely with scattered hairs on the margins, often persistent; flower clusters with 615 flowers, spirally arranged on inflorescence branches; bracteoles broadly ovate, 46 by 34 mm, margin irregularly deeply dentate with spreading narrow unequal teeth, apex broadly acute, both surfaces covered in minute silvery to ferrugineus hairs. Staminate flowers c. 0.7 mm long, shortly pedicellate; sepals 3, free, densely pubescent, stamens 23; anthers 4-locular. Pistillate inflorescences erect narrow panicles usually not extending above the leaves, 1228 cm long, 3 axis orders, basal unbranched axis slightly flattened and less than 1/2 (usually less than 1/3) of the total inflorescence length, basal axes glabrous, distal axes silvery to ferrugineus pubescent; bracts ovate, 510 mm long, margin entire, glabrous, caducous. Pistillate flowers c. 1.5 mm long; calyx urceolate, c. 1 mm long, densely ferrugineus pubescent; ovary 2-carpellate; styles c. 1 mm long, free, persistent. Fruits bilocular, 45 by 56 mm, turning red at maturity, locules subglobose, without processes, covered in yellowish-green, sticky exudate; pedicel slender, 58 mm long, with ferrugineus hairs. Seeds c. 3 mm in diam., lenticular, black, shallowly pitted, encased in a fleshy violet aril.
Distribution Borneo (Brunei, Sarawak).
Habitat & Ecology Macaranga rufescens is a large early successional tree. It is a tree of lowland mixed dipterocarp forest, and appears to be more common on more fertile soils, such as clay-rich ultisols. On good soils this species can become one of the largest species in the genus in Borneo, reaching above 30 m in height and 50 cm dbh. However, it may also be found flowering at c. 10 m in height in secondary successional forest.
Note Macaranga rufescens is most closely related to M. hosei. It differs most obviously from M. hosei in having short basal axes on both staminate and pistillate inflorescences, dense minute ferrugineus hairs on the adaxial surface of young leaves, few or no erect hairs on the lower surface of mature leaves, an absence of conical nectaries along the base of the leaf margin, and considerably larger leaves, stipules and twigs. Further differences between these two species are described under M. hosei.
Macaranga rufibarbis Warb., Bot. Jahrb. 16 (1893) 21; K.Schum. & Lauterb., Fl. Schutzgeb. Sόdsee (1900) 397 in part; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 373; L.M.Perry, J. Arnold Arb. 34 (1953) 211; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 156; Gen. Macaranga Prodr. (2008) 230. Type: Hellwig 308 (not seen), Papua New Guinea, Morobe Prov., Finisterre Mountains.
var. rufibarbis
var. rufibarbis: Whitmore, Gen. Macaranga Prodr. (2008) 230. Type: as the species.
Small trees, to 5m. Twigs slender, 3 mm diam., terete, at first finely rufous pubescent, becoming glabrous, ringed by stipule scars. Stipules elliptic, 20 by 3 mm, apex acuminate, chestnut brown, papery, variously rufous hirsute on outer side, erect, subpersistent. Leaves: petioles 25.5 cm, slender, 1(2) mm diam., finely rufous pubescent or glabrous, kneed at top; blades deltoid, size very variable, 1020 by 716 cm, thinly to thickly leathery, base very broadly rounded to truncate, 1 mm peltate or cordulate, with 2 large glands near petiole insertion, 3-nerved, margin slightly recurved especially at base and with close small protruding glandular teeth, apex acuminate to subcaudate, drying greyish above, rufous below, usually at least some leaves at least partially bullate, glabrous above, below with dense dark to black granular glands and variously tomentose from dense on all nerves to sparse, sometimes hairy only near petiole insertion, rarely glabrous, hairs often tufted. Staminate inflorescences diffuse racemes or 3 axis-order panicles, 415 cm long, rufous tomentose; branches normal to ascending; peduncle 23 cm; main axis 1.52 mm diam., terete, lowest branches opposite; bracts variable, 36 by 13 mm, papery, almost glabrous to hirsute, soon caducous; all branches arising from a flower cluster, to 5 cm long; compound flower clusters evenly 510 mm spaced, consisting of a tight mass of a few clusters, each subtended by a squarish thick spreading bracteole; latter 5 by 3 mm or less, apex blunt, margin distantly shallowly irregularly toothed, glabrous within without patellar glands, hirsute without, smaller than cluster. Staminate flowers c.3 per cluster, hirsute; stamens c. 11, anthers 4-locular. Infructescences spikes to 11 cm, with an apical cluster of 13 fruits, subtending bracts not seen; axis 1.5 mm diam., terete, striate, usually almost glabrous, with at halfway bracts or their scars, latter elliptic, 1 cm, pubescent, caducous. Fruits globose, 6 mm, woody, densely invested with flattened soft rufous hirsute 5 mm spines; pedicel stout, 310 mm, tomentose; calyx persistent; style apical. Seeds round, coarsely verrucose, with sarcotesta.
Distribution Throughout New Guinea
Habitat & Ecology Secondary montane forests, 14001900 m.
var. campestris Whitmore
var. campestris Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)156; Gen. Macaranga Prodr. (2008) 231. Type: Streimann & Lelen 34140 (holo K; iso SING), Papua New Guinea, Western Distr., Kiunga.
Differs solely in the small 5 by 1.5 mm stipules and lowland habitat.
Distribution New Guinea (Western district; known from type only).
var. tenella (Pax & K.Hoffm.) L.M.Perry
var. tenella (Pax & K.Hoffm.) L.M.Perry, J. Arnold Arb. 34 (1953) 212; Whitmore, Gen. Macaranga Prodr. (2008) 231. Macaranga tenella Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 379. Syntypes: Schlechter 18901 (not seen), Papua New Guinea, Madang Prov., Bismarckgebirge; Schlecher 19101 (K, P), Papua New Guinea, Morobe Prov., Finisterre-Gebirge; Schultze 283 (not seen), Papua New Guinea, Sepik Region, August River.
Differs solely but distinctly in its leaves elliptic, 917 by 26 cm.
Distribution East New Guinea (Finisterre Range). Only twice collected.
Habitat & Ecology Mountains at 1300 m.
Macaranga salicifolia Airy Shaw, Kew Bull. 23 (1969) 108; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 157; Gen. Macaranga Prodr. (2008) 232. Type: P. van Royen & Sleumer 8080 (holo K; iso L, P, SING), Indonesia, Papua, Vogelkop, Nettoti Range, Wekari River, path to Saokorem.
Tall tree, 27 m; bole 20 m, 50cm diam. Twigs slender, terete, 3 mm diam., at first tawny tomentose, soon glabrous, blackish brown, with stipule scars. Stipules united, apparently single, forming a long slender elliptic terminal bud, 9 by 0.8 cm, grey-brown sericeous. Leaves: petioles 1 cm, slender, terete, sericeous, kneed at top; blades elliptic, 13 by 1.5 cm, leathery, base long tapering but minutely rounded at petiole and with 15 near-basal glands on lamina, margins revolute, apex long-tapering, drying greyish, below with brown main nerves, shagreened and glabrous above, below midrib with sericeous hairs and dark granular glandular. Staminate unknown. Infructescences slender 7 cm axillary spikes; axis slender, 0.8 mm, slightly flattened, sericeous and sparsely granular glandular; bracteoles 23, dissimilar: one spathe-like, 6 by 12 mm, apex obtuse, adpressed pilose and granular glandular outside with a terminal gland-bearing limb, plus 12 others subulate, 35 mm, dorsally pilose. Fruits apical, solitary, globose, 10 mm (? immature), densely invested with slender 3 mm hairy spines; pedicel 1 cm; calyx bladder-like, narrowed into a terminal collar, sparsely tomentose, papery, dark brown; styles 2, apical, slender, 15 mm, papillose, caducous, slender, initially tightly enclosed in calyx. Seeds unknown.
Distribution West New Guinea (known from type only).
Habitat & Ecology Montane forest, 1550 m.
Note A big, highly distinctive tree. As Airy Shaw (1969) pointed out, the leaves have a pair of (faint) longitudinal folds reminiscent of Erythroxylum.
Macaranga salomonensis L.M.Perry, J. Arnold Arb. 34 (1953) 210; Airy Shaw, Kew Bull. 23 (1969) 98, 99 passim; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 157; Gen. Macaranga Prodr. (2008) 232. Type: Brass 3327 (holo A; iso L), Solomon Islands, Santa Isabel Island.
Small trees to 10 m. Twigs slender, 34 mm diam., coarsely angled, at first pubescent, soon glabrous. Stipules ovate-oblong, 10 by 8 mm, apex acute, dark to blackish brown, thinly papery, crumpled, pubescent to glabrous, spreading to reflexed, usually persistent. Leaves: petioles 18 cm, slender, 1.5 mm diam., slightly flattened, glabrous, sometimes drying weakly constricted at base; blades ovate, c. 17 by 14 cm, papery, base broadly rounded, 4 mm peltate, margin repand with small protruding inconspicuous glandular teeth, apex abruptly acuminate, 8 mm, drying khaki, grey-glaucous below with dark nerves, above glabrous, below (innovations densely tomentose) weakly pubescent to glabrous with tiny golden granular glands, spider's web nervation, tertiaries scalariform. Staminate inflorescences 617 cm sparsely branched racemes with 27 cm glabrous peduncle; branches 1.54 cm, tomentellose, becoming glabrous; bracteoles concave, semiorbicular, 3 mm, rather thick, adaxially with more or less confluent patellar glands, recurved; axis zigzagging between distant flowers clusters. Staminate flowers c. 25 per cluster; pedicel 1 mm, pubescent; stamens 58, anthers 4-locular. Infructescences 9 cm spikes with a single terminal fruit at first subtended by 1 cm diam. roundish dentate leafy bracts; axis flattened, 2 mm across, glabrous. Fruits weakly bilobed, 10 by 15 mm, blackish, woody, golden granular glandular with dense to sparse 5 mm slender soft sinuous spines; pedicel stout, 5 mm, glabrous; calyx caducous. Seeds 8 mm diam., rugulose.
Distribution Papua New Guinea (Northern Division, Bougainville), Solomon Islands (Kolombangara, Santa Ysabel, Guadalcanal).
Habitat & Ecology Primary or regrowth forest. Lowlands to 500 m,
Note Staminate parts described partly from Perry (1953), on whose authority patellar glands are described on the bracteoles unique in the group.
Macaranga sandsii Whitmore, Gen. Macaranga Prodr. (2008) 234. Type: Sands 368 (holo K; iso L), Sulawesi, Enrekang, Latimojong Mountains.
Small tree. Twigs somewhat angular, coarsely striate, fairly slender, 4 mm diam., dark brown, glabrous. Stipules elliptic, to 50 by 5 mm, broadest at base and tapering into acuminate tip, chestnut brown, papery, glabrous, forming a conspicuous elliptic terminal bud, becoming reflexed, very soon caducous, leaving conspicuous scars. Leaves: petioles 8 cm, slender, flattish, striate, glabrous, with weak glaucous apical knee; blades ovate-elliptic, 1518 by 78 cm, thinly leathery, base rounded, 2 cm peltate, margin with 1 cm distant protruding conspicuous dark glands, apex narrowing suddenly to a 3 cm acuminate tip, drying dark brown above, ochreous below, undersurface closely golden granular glandular and with single white spreading hairs on main nerves, 5 secondary nerves radiating from petiole insertion otherwise pinnately nerved. Staminate inflorescences (young unexpanded) subterminal, to 10 cm, peduncle to 5 cm, flattened, dark brown, slightly glaucous, with opposite branches and often axillary branches subtended by a caducous bract (only the scar seen), these branches 1 cm terminated by an ovate to oblong-elliptic, dark brown, papery, glabrous, 12 cm caducous bract enclosing a spike to 2 cm bearing spaced 3 mm, c. 10 flowered clusters, themselves subtended by a persistent, blackish brown, glabrous, linear 3 mm bracteole widened to an adaxial patellar gland bearing tip. Stamiante flowers: stamens with anthers 4-locular. Pistillate unknown.
Distribution Sulawesi (known from type only).
Habitat & Ecology Montane oak-Podocarpus forest, 1650 m.
Notes 1. This is a highly distinctive species, known so far from only one collection. Notable are the big elliptic stipules, forming a conspicuous terminal bud, peltate leaves ochreous below and with conspicuous protruding marginal glands, and staminate inflorescence construction. The inflorescences are mostly young, but one contains open staminate flowers, the conspicuous bracts drop off as it expands.
2. Most remarkable of all is that another new species, M. serratifolia, also with leaves ochreous below, grew adjacent to it.
3. I place this in the Angustifolia group, with which the glandular staminate bracteoles is its most discordant character.
Macaranga sarcocarpa Airy Shaw, Kew Bull. 23 (1969) 88; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 152; Gen. Macaranga Prodr. (2008) 235. Type: Haviland & Hose 3663K (holo K), Borneo, Sarawak, Kuching.
Small tree(?). Twigs stout, 8 mm diam., terete with prominent encircling stipule scars, tawny tomentose. Stipules ovate, 12 by 8 mm, apex acute, dark brown, papery, outside tawny pubescent, inside glabrous, erect, soon caducous. Leaves: petioles 818 cm, stout, 3 mm diam., terete, tawny tomentose; blades orbicular-ovate, c. 17 by 14 cm, thickly leathery, base broadly rounded, 47 cm peltate, margin with distant protruding glandular teeth, apex acute, 1 cm acuminate, or obtuse, drying tawny brown, all nerves prominently raised below, above bullate, below tomentose with tufted white hairs, densely granular glandular. Staminate unknown. Infructescences short spikes to 5 cm, axis stout, 3 mm diam., slightly flattened, striate, tawny pubescent; peduncle 1 cm. Fruits mostly apical, crowded, depressed globose, 15 mm diam., black, closely granular glandular, bilobed, fleshy; pedicel very stout, 410 mm long, furfuraceous; sepals 4 mm, furfuraceous; stigmas 2 mm, apical, persistent, filiform. Seeds globose, 45 mm diam., with sarcotesta.
Distribution Borneo (Sarawak, near Kuching).
Note The leathery, bullate leaves, reminiscent below of Victoria amazonica, and the big fleshy fruits are highly distinctive.
Macaranga serratifolia Whitmore, Gen. Macaranga Prodr. (2008) 238. Type: Sands 366 (holo K; iso L), Sulawesi, South, Enrekang, Latimojong Mountains.
Small tree. Twigs slender, 12 mm diam., round, shallowly coarsely striate, dark brown, weakly pubescent to glabrous. Stipules narrowly elliptic, 4 by 1 mm, brown, papery, glabrous, spreading, most caducous. Leaves: petioles 1.5 3 cm, slender, round, terete, glabrous; blades ovate-elliptic, 512 by 1.54 cm, thinly leathery, base (broadly) cuneate, sometimes 0.5 mm peltate, usually with 2 inconspicuous glands at petiole insertion, margin shallowly crenate, or deeply serrate with dark marginal glands, apex 2 cm acuminate to caudate, drying grey-green above and sometimes ochreous below, underside (closely) golden granular glandular and with main nerves sometimes slightly finely pubescent. Staminate inflorescences short slender racemes or spikes to 6 cm, axes slender, slightly flattened, glabrous, secondaries spreading to 1 cm, subtended by minute persistent bracts; flower clusters spaced to 1 cm, to 12-flowered; bracteoles spreading, elliptic, 3 by 1 mm, glabrous, with 13 adaxial patellar glands. Staminate flowers 0.3 mm, shortly pedicellate; stamens 2 or 6, anthers 4-locular. Pistillate inflorescences as staminate, spikes 25 cm. Fruits distant, solitary, (young) globose, somewhat flattened, weakly bilobed, 3 mm across, sparsely granular glandular, sessile; calyx subpersistent; styles 2, persistent, recurved, 1 mm, apical. Seeds unknown.
Distribution Sulawesi.
Habitat & Ecology Mountains (primary?) forests, 11001650 m.
Notes 1. The two specimens, both from primary montane forest in Sulawesi are not identical. The type has small strongly serrate leaves and 2 stamens per flower. Musser 974 has larger, crenate leaves and 6 stamens, despite which, I believe it is conspecific.
2. See note at M. sandsii.
Macaranga setosa Gage, Rec. Bot. Surv. India 9 (1922) 244; Ridl., Fl. Malay. Penins. 3 (1924) 304; Airy Shaw, Kew Bull. 19 (1965) 317; Whitmore, Tree Fl. Malaya 2 (1973 112; Gen. Macaranga Prodr. (2008) 239. Type: Lake & Kelsall s.n. (iso SING), Peninsular Malaysia, Johore, Sungai Ulubrong.
Small tree. Twigs 3 mm, angled, caducous strigose and (densely) pubescent. Stipules subulate, 5 mm, erect, subpersistent, strigose. Leaves: petioles 47 cm, slender, 1.5 mm diam., terete, pubescent with sparse spreading white caducous hairs, kneed at both ends; blades ovate, c. 10 by 5 cm, thinly leathery, base broadly rounded, minutely cordulate, the lobes overlapping and without glands, apex acuminate, drying greyish above, fawn below, below the main nerves sparsely pubescent, densely granular glandular. Staminate inflorescences 12 cm spikes; peduncle short; axis angular, striate, pubescent and granular glandular; flower clusters diffusely grouped; bracteoles erect, ovate, 2 by 1 mm, apex acuminate, outside sparsely strigose, both sides sparsely granular glandular. Staminate flowers c. 12 per cluster, sessile; sepals granular glandular and sparsely strigose; stamens 1012, anthers 4-locular. Infructescences axis 46 cm, angled, ridged, strigose and finely pubescent with a pair of apical orbicular bracts, c. 2 by 3 cm, shortly caudate-acuminate, strigose and finely pubescent and with sparse granular glands, pinnately nerved. Fruits not seen.
Distribution India (Anambas Islands), Peninsular Malaysia (S.W. Pahang, Johore).
Habitat & Ecology Primary lowland forest.
Note Still only known from three collections. The flower description is taken from Gage (1922).
Macaranga similis Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv add. vi (1919) 29; L.M.Perry, J. Arnold Arb. 34 (1953) 225; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 157; Gen. Macaranga Prodr. (2008) 239. Type: Ledermann 12622a (not seen), Papua New Guinea, Sepik Region, Felsspitze.
Macaranga brassii Mansfeld, J.Arnold Arbor. 10 (1929) 78, 232.
Small trees, 510 m. Twigs fairly stout, 4 mm diam., weakly squarish to almost terete, densely pale velvety tomentose, hairs simple or tufted. Stipules oblong, 15 by 6 mm, broad rounded base, narrowing upwards to acute or acuminate apex, often carinate, outside pale velvety tomentose, inside dark brown almost glabrous, papery, erect, persistent. Leaves: petioles c. 12 cm, terete, 2 mm diam., tomentose as twigs; blades deltoid, broadest near base, 1324 by 1318 cm, very rarely with small broad lateral cusps, thinly leathery, base truncate but with 1cm deep narrow cordate sinus, the lobes almost touching, usually several small inconspicuous glands or nerves near petiole insertion, margin weakly repand with inconspicuous distant protruding glands, apex 2 cm acuminate rarely caudate, drying warm khaki brown, densely velvety below from all nerves with white simple and tufted hairs dark brown granular glandular, above velvety to the touch from lamina with sparse and main nerves with dense hairs, spider's-web nervation, palmate with scalariform tertiaries. Staminate inflorescences diffuse racemes or panicles with 3 axis orders, to 22 cm long, tomentose as twigs,; branches divaricate; peduncle 47 cm; main axis fairly slender, 1.5 mm diam., terete, branches to 5 cm, most arising from a flower cluster, thread-like, lowermost branches subopposite; bracts ovate, 3 by 1 mm, apex acute, tomentose outside, glabrous inside (like stipules), persistent; flower clusters 3 mm diam., 35 mm evenly spaced like beads on a thread, to contiguous; bracteoles recurved, as long as clusters, triangular, narrowing above middle, 3 by 2 mm, apex acute, margin entire, abaxial surface tomentose, adaxial glabrous with 24 patellar glands near margin towards apex. Staminate flowers c.6 per cluster, 1 mm with a few granular glands; pedicel 1 mm; stamens 3, anthers 4-locular. Infructescences 10 cm spikes to 25 cm sparsely 3 cm branched racemes with crowded terminal 2 by 2 cm clusters of leafy bracts subtending fruits; main axis flattened, 2 mm diam., sinuous, velvety as twigs; bracts subtending branches as staminate but to 5 mm; bracts leaf-like, ovate, to 2 by 1 cm, apex acuminate, subsessile, base broadly rounded, margin finely serrate, thinly leathery, abaxial surface velvety, adaxial with scattered hairs and dark granular glands, pinnately nerved. Fruits globose, 4 mm, thinly woody, with close black granular glands, at first with 3 mm pubescent spines, becoming unarmed; pedicel 3 mm, pubescent; calyx persistent, adherent; style lateral, 1015 mm, thread-like, erect, with short close comb-like hairs along outer side, caducous. Seeds round, 3 mm, coarsely verrucose, with sarcotesta.
Distribution East New Guinea (common in Milne Bay District), Solomon Islands.
Habitat & Ecology Gregarious in lowland secondary forest to 500 m.
Notes 1. Easily recognised from its leaves, velvety on both surfaces with some hairs tufted. Neither Perry (1953) nor I (Whitmore 1980) have seen the single sheet from which the species was described, but the staminate bracteole matches, not least in its unusual shape.
2. Macaranga dioica and M. galorei (see there) are close.
Macaranga sinensis Baill. ex Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1001; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 351; Gagnep., Bull. Soc. Bot. France 69 (1923) 705; Merr., Philipp. J. Sci. 60 (1936) 30; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 35; Chiu, Guihaia 2 (1982) 148, in key; Whitmore, Gen. Macaranga Prodr. (2008) 240. [Mappa sinensis Baill., Ιtude Euphorb. (1858) 430, nomen subnudum]. Type: Callιry 9 (holo P), China.
Macaranga dipterocarpifolia Merr., Philipp. J. Sc. 1 Suppl. (1906) 205; Enum. Philipp. Flow. Pl. (1923) 441; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 338; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 34.
Small trees, 7 m. Twigs slender, rounded, glabrous. Stipules ovate-oblong, 14 by 3 mm, apex acute, papery, glabrous, somewhat persistent, spreading to recurved. Leaves: petioles 410 cm, slender, terete, glabrous; blades broadly ovate, 815 by 411 cm, thinly leathery, base very broadly truncate, usually with 2 glands, sometimes minutely peltate, margin usually weakly crenate, apex acute to acuminate, glabrous with very inconspicuous granular glands below. Inflorescences somewhat crowded, racemes to 14 cm, 3 orders of slender glabrous branches sometimes with axillary branches, secondaries to 8 cm; flower clusters 1 cm spaced, tiny, 1 mm across; bracteoles linear, 3 mm, spreading well beyond cluster, soon caducous, broadening to acute tip, bearing 12 patellar glands. Staminate flowers c. 10 per cluster, 0.08 mm diam.; pedicel long; calyx divided to base; stamens 56, anthers 4-locular. Pistillate flowers single or paired, pedicellate; calyx shallowly lobed. Fruits crowded, solitary or paired, small round, slightly bilobed, 3 mm across, smooth, glabrous or finely granular glandular, thinly woody; pedicels slender, 3 mm; calyx small, persistent; styles persistent, recurved, short, flattish, terminal. Seeds 2, round, pitted.
Distribution China, Taiwan, Philippines (Luzon, Ilocos Norte, Batan, Leyte, Mindanao), Sulawesi (Menado, once collected).
Habitat & Ecology 4001300 m (Philippines).
Macaranga spathicalyx Whitmore & S.J.Davies, Harvard Pap. Bot. 6 (2001) 367; Whitmore, Gen. Macaranga Prodr. (2008) 241. Type: Gibot SAN 79573 (holo K; iso L, SAR, SING), Borneo, Sabah, Mt. Kinabalu National Park.
Trees to 17 m. Twigs 5 mm diam., angled at tip, terete behind, blackish brown, glabrous. Stipules ovate, strongly asymmetric, 815 by 48 mm, apex acute, with a central nerve, dark brown, papery, furfuraceous but soon glabrous, erect, mostly soon caducous. Leaves: petioles 512(20) cm, slender, terete, slightly ridged, glabrous, not kneed; blades ovate, 1022 by 816 cm, papery, base broadly rounded to subtruncate, (2)520 mm peltate, usually with 2 large 3 mm conspicuous glands near petiole insertion, margin entire sometimes very weakly crenate, apex acute, drying grey green above dark brown below, glabrous, closely minutely granular glandular. Staminate inflorescences amongst and behind leaves, 613 by 48 cm panicles, with 3 axis orders; peduncle under 1 cm; main axis terete or slightly flattened; branches alternate; bracts 2, lateral, 35 by 1 mm, asymmetric, stipuliform structures plus a central elliptic1020 by 38 mm acute leaf-like bract with margin toothed and 36 prominent round flat abaxial glands, glabrous, caducous; flower clusters spiral, evenly spaced on ultimate branches; bracteoles triangular, c. 4 by 3 mm, apex acute, margin entire, glabrous. Staminate flowers 715 per cluster, c. 4 mm long; pedicel 11.5 mm, persistent after fall of flowers; sepals 3, free, reflexed after anthesis, glabrous; stamens 812, anthers 4-locular. Pistillate inflorescences amongst and behind leaves, crowded, entangled, diffuse panicles, 814 cm long, 3 axis orders, glabrous; main branches divaricate to 4 cm; peduncle c. 1 cm; main axis stout, 1.5 mm diam., terete or slightly flattened; lower branches alternate, sometimes with short axillary branches; bracts soon caducous, lateral stipuliform structures as staminate, central bract similar to but narrower than staminate, 820 by 14 mm; ultimate branches thread-like, sometimes arising from flower clusters; bracteoles narrowly elliptic, variable, 38 by 11.5 mm, apex long caudate, dark brown, papery, curling when dry, sparsely furfurescent, the larger with 12 patellar glands, with a central nerve, erect, persistent, subtending clusters of c. 3 flowers, c. 1 cm apart, evenly spaced. Pistillate flowers c. 10 mm long; calyx spathe-like, 34 mm long, 1.5 mm wide, split almost to base on one side, with 34 short sharp ribbed teeth extending beyond apex, glabrous, persistent. Young fruits enclosed by the spathe-like dark brown 3 mm papery calyx, with twin apical brittle 69 mm erect styles fused at base, coarsely papillose, caducous. Mature fruits 34 by 46 mm with 2 globular loculi, closely yellowish granular glandular, with 410 soft curly slender 36 mm caducous spines, often asymmetric on the 1020(25) mm slender glabrous pedicel. Seeds spheroidal to ellipsoidal, 3 mm diam., with shallow grooves and pits, enclosed in a fleshy black sarcotesta.
Distribution Borneo (Sarawak, Sabah).
Habitat & Ecology Lower montane rain forest, 16001800 m.
Notes 1. Very distinctive in its big ovate asymmetric nerved stipules, peltate leaves and spathaceous calyx persisting into young fruit.
2. Creagh s.n. [May 1895] (K!) from Pulau Gaya is a very young staminate. It matches this species but differs in having the adaxial surface of the leaves finely pubescent on nerves and lamina, and in its lowland station.
Macaranga stellimontium Whitmore, Kew Bull. 34 (1979) 604; in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 157; Gen. Macaranga Prodr. (2008) 244. Type: Vinas & Wiakabu 59632 (holo K; iso L), Papua New Guinea, West Sepik Dist., Telefomin Subdist., Star Mountains near Busilmin.
Small tree, 6 m. Twigs 34 mm diam., squarish, golden strigose at first densely. Stipules broadly triangular, 15 by 6 mm, base truncate wider than twig, apex acute, papery, tomentose with dense adpressed long golden hairs, erect, persistent. Leaves: petioles 27 cm, slender, 2 mm diam., terete, golden pubescent, and at first sparsely strigose, kneed at both ends; blades broadly ovate, c. 12 by 9 cm, thinly leathery, base broadly truncate, minutely cordulate at petiole insertion, margin with small distant glandular teeth, apex acuminate, drying chocolate brown, very finely bullate (appearing shagreened), velvety on both surfaces from rather sparse white hairs, above with sparse granular glands, with 3 main and several minor basal nerves. Staminate inflorescences narrow sparsely branched 614 cm finely velvety racemes; branches slender, sinuous, 3 cm, ascending, arising from a flower cluster, lowermost pair opposite with 5 mm axillary branches; peduncle long, 49 cm; main axis flattened, 1.5 mm diam.; flower clusters small, in 3mm spaced groups; bracteoles spathulate, 2 by 1.2 mm, slightly broadening to trapezoid acute head, fleshy, outside pubescent, inside with 45 patellar glands, spreading, persistent, longer than cluster. Staminate flowers 16 per cluster, obconical, sessile; sepals glabrous, tipped by a dark gland; stamens 2, anthers 4-locular. Pistillate unknown.
Distribution East New Guinea (known from type only).
Habitat & Ecology Secondary montane forest, 1500 m.
Note Very distinctive.
Macaranga stenophylla Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 371; L.M.Perry, J. Arnold Arb. 34 (1953) 211; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 162; Gen. Macaranga Prodr. (2008) 245. Type: Schlechter 17724 (not seen), Papua New Guinea, Madang Prov., Kani Gebirge.
Twigs slender, at first rusty brown tomentose. Stipules not seen, soon caducous. Leaves: petioles 4.57 cm, tomentose; blades elliptic-ovate, oblong or oblong-lanceolate, 1218 by 67 cm, papery, base rounded, 710 mm peltate, 3-nerved, with several glands between base and petiole insertion, apex acuminate, above glabrous, below reticulate and sparsely puberulous on nerves, granular glandular. Staminate inflorescences rusty brown tomentose panicles; clusters many-flowered; bracteoles triangular with apex elliptic, tomentose with adaxial patellar glands. Staminate flowers with turbinate calyx, granular-glandular and tomentellose; stamens 1516, anthers 4-locular. Pistillate unknown.
Distribution East New Guinea (known from the type only).
Habitat & Ecology 1000 m.
Notes 1. The type collection is from a staminate plant with young flowers; not seen by me or Perry (1953). The description is paraphrased from Perry's (1953) translation, amended against the original Latin.
2. This keys out to the Brunneo-floccosa group but not to any known species therein. The leaves 710 mm peltate plus patellar glandular staminate bracteoles make it distinctive, despite the rather sketchy description. In the absence of data it is entered in both parts of the key, as leaves both bullate and non bullate.
Macaranga sterrophylla L.M.Perry, J. Arnold Arb. 34 (1953) 231; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 158; Gen. Macaranga Prodr. (2008) 245. Type: Brass 4560 (holo A; iso BO), Papua New Guinea, Central Prov., Murray Pass, Wharton Range.
Trees, 815 m. Twigs 3 mm diam., terete, densely rusty pubescent or tomentose. Stipules elliptic, 9 by 3 mm, apex acute, without densely rusty pubescent, glabrous within, papery, erect, soon caducous. Leaves: petioles c. 2.5 cm, slender, 11.5 mm diam., terete, pubescent to furfuraceous, kneed at both ends; blades ovate to elliptic, 49 by 34 cm, leathery, base rounded or cuneate, to 1 mm peltate or cordulate, with a few small inconspicuous glands near petiole insertion and 3 main plus several minor nerves, margin (weakly) repand, apex acute to 1 cm acuminate, drying pale brown to khaki, below and above initially densely rusty floccose, later above glabrous shiny with reticulum fine and raised giving shagreened appearance (not bullate), and below tomentose just on midrib, sparsely to densely golden or dark granular glandular, reticulum paler than lamina on upper and also sometimes on lower surface, secondaries paler or darker. Staminate inflorescences rusty tomentose 4 cm racemes with short 1 cm erect slender branches; peduncle 1.2 cm; main axis <1 mm diam., terete; branches alternate; bracts linear, 10 mm, erect, papery tomentose outside, caducous; flower clusters irregularly spaced; bracteoles narrowly trapezoid, 5 by 1 mm, broadest near acuminate tip, narrowing to a basal stalk, thick, with 2 rows of small adaxial patellar glands, much larger than cluster, spreading, persistent. Staminate flowers in clusters of 4, sessile; sepals 2; stamens 3, anthers 4-locular. Infructescences rusty pubescent 310 cm spikes or sparsely short-branched racemes. Fruits solitary, unilocular, ellipsoid, 7 by 4 mm, leathery, evanescent rusty pubescent, rarely sparsely tuberculate, inside wall tomentose; calyx big, sessile, persistent; stigma short, 1 mm, subapical, persistent. Seeds roundish, 5 mm, densely verrucose, with sarcotesta.
Distribution Southeast New Guinea.
Habitat & Ecology Primary montane forests, 20002900 m.
Note The leaf reticulations pale on both surfaces are very distinctive, as are the staminate bracteoles. Hairs are present on the inside ovary wall also in M. brunneo-floccosa and M. carrii.
Macaranga stonei Whitmore, Gen. Macaranga Prodr. (2008) 247. Type: Soejarto, Fernando & Fernando 7957 (holo L), Philippines, Luzon, Aurora Prov., road San Luis Cabanatuan.
Treelet 8 m. Twigs 3 mm diam., slightly angular, glabrous. Stipules oblong-elliptic, 20 by 4 mm, tapering to a long acumen, with a central nerve, dark brown, papery, margins recurved, glabrous, recurved, subpersistent. Leaves: petioles 10 cm, slender, 2 mm diam., terete weakly ridged, glabrous, weakly kneed at top; blades deltoid, c. 19 by 13 cm, papery, base broadly truncate, 3 mm peltate, margin coarsely closely crenate finely recurved, apex acuminate, 2 cm, drying dark brown, glabrous, abaxial surface with very dense close dark granular glands, secondary nerves c.13 pairs, straight and parallel, basal pair plus other minor ones from petiole insertion. Staminate inflorescences open delicate glabrous panicles 8 by 4 cm, with 3 axis orders, branches ascending to divaricate; peduncle 2 cm, terete, slender, 1.5 mm diam.; lowest branch with an axillary branch; bracts elliptic, 4 by 1.5 mm, apex acuminate, entire, papery, glabrous, spreading, caducous; ultimate branches flattish thread-like, sometimes slightly zigzag between 35 mm spaced small flower clusters; bracteoles to 5 mm long, spoon-like with a narrow stalk and an enlarged 1 mm diam. gland-bearing roundish apex, entire, spreading, much longer than cluster, soon caducous or breaking off. Staminate flowers 8 per cluster, glabrous, shortly pedicellate; sepals free, tipped by granular glands; stamens 4, anthers 4-locular. Pistillate unknown.
Distribution Philippines (known from type only).
Habitat & Ecology Logged lowland rain forest, deep shade, 200300 m.
Notes 1. Named for Benjamin J. Stone (19331994), who devoted his last few years to study of the Philippines' flora.
2. This species has no obvious affinity with any of the groups I recognise.
Macaranga strigosa Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv add. vi (1919) 30; L.M.Perry, J. Arnold Arb. 34 (1953) 214; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 158; Gen. Macaranga Prodr. (2008) 248. Type: Ledermann 10939 (not seen), Papua New Guinea, Sepik Region, Hunsteinspitze.
Macaranga bullata Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv add. vi (1919) 31; L.M.Perry, J. Arnold Arb. 34 (1953) 215; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 161.
Macaranga glandulifera L.M.Perry, J. Arnold Arb. 34 (1953) 218; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 161.
var. strigosa
var. strigosa: Whitmore, Gen. Macaranga Prodr. (2008) 248. Type: as the species.
Shrubs, 56 m. Twigs stout, terete, with spreading rigid whitish hairs drying fulvous, tardily glabrescent. Stipules triangular-ovate, 45 by 2.53 mm, apex acute, rigid, keeled, adpressed villous, erect, soon caducous. Leaves: petioles 312 cm, slender, terete, with dense to sparse long pale hairs; blades deltoid, 917 by 716 cm, thinly leathery, base broadly truncate, very broadly shallowly cordate plus at petiole 3 mm cordulate with overlapping or touching lobes, several large glands on lobes near petiole insertion, margin with small 5 mm distant protruding glandular teeth, apex acuminate, drying dark chocolate brown, finely weakly bullate, above with a few long hairs on main nerves, below all nerves prominent and with dense solitary and tufted white spreading hairs on nerves and black granular glandular, with 3 main basal nerves. Staminate inflorescences villous 1114 cm panicles; peduncle 79 cm; bracts 3, elliptic, 1025 mm, scarlet, stipulate, about equal in length to inflorescence branches; bracteoles 3 mm long, 3-lobed, pilose outside, longer central lobe with 4 patellar glands. Staminate flowers in clusters of 1215; sepals 3, scarcely granular glandular; stamens 3(4), anthers 34-locular. Pistillate inflorescences racemes 618 cm long, glabrescent, with a few small branches to 3 cm long bearing 12 apical flowers, with small sterile bracts above the middle, oblong, 7 mm long, narrowed at base, outside sparsely pilose and inside granular glandular, with sometimes confluent patellar glands. Pistillate flowers: ovary 2-locular, densely covered with processes; styles 7 mm, plumose. Fruits (immature) globose, to 7 mm diam. including dense soft flattened 3 mm finely pubescent spines.
Distribution New Guinea.
Notes 1. I cannot see any difference between this species and Perry's (1953) M. glandulifera which I therefore reduce. The description is partly a paraphrased translation of the Latin originals.
2. Macaranga bullata was described tersely from one collection at B (now destroyed) from Mt. Schrader and contrasted with this in leaves 1 cm peltate, more pyramidal and open male inflorescences, no [i.e., caducous] staminate bracts, granular glandular calyx and all stamens 4-locular. I consider all these characters within the probable range of variation of a single species and have little doubt that if an isotype is ever found it will prove to be M. strigosa.
var. carrii L.M.Perry
var. carrii L.M.Perry, J. Arnold Arb. 34 (1953) 214; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)158; Gen. Macaranga Prodr. (2008) 248. Type: Carr 13231 (holo A; iso BM, K, L, SING), Papua New Guinea, Central Prov., Boridi.
Small trees, 46 m. Stipules ovate, 815 mm, apex acuminate, concave, erect, subpersistent. Leaves: blades deltoid-ovate, 820 by 614 cm, occasionally lobulate, base subtruncate sometimes cordulate sometimes with 2 glands and with 35 basal nerves, apex 2.5 cm, at maturity, above with main nerves pubescent, below sparsely granular glandular and sometimes sparsely pilose, not bullate. Staminate inflorescences with bracteoles 2.5 mm long. Staminate flowers 0.6 mm with 2 sepals; stamens 23, 4-locular. Pistillate inflorescences to 23 cm; bracts apical, long triangular-ovate, 4.58 mm, narrowed to the base, patellar glandular within. Pistillate flowers urceolate, 1.52 mm long, glabrate; styles basal to ovary, 2, connate, 9 mm, papillose and shortly plumose. Fruits 8 by 10 mm (almost mature), spines 1.5 mm, more or less caducous.
Distribution East New Guinea (mountains of the southeast peninsula from the Murray Pass to Myola and Milne Bay district).
Habitat & Ecology Landslips, regrowth forest and primary lower and upper montane rain forest; 14002400 m
Notes 1. Perry (1953) says var. carrii differs in somewhat larger stipules, smaller staminate bracteoles (though she said bracts) and lack of bracts along the peduncle of the pistillate inflorescence, as well as in leaf size, shape, vestiture and absence of bullae.
2. LAE 54958 is hermaphrodite.
Macaranga strigosissima Airy Shaw, Kew Bull. 19 (1965) 320; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 155; Gen. Macaranga Prodr. (2008) 249. Type: Anderson 14705 (holo K; iso L, SING), Bonreo, Sarawak, Kapit Distr., Bukit Raya, Pelagus.
Macaranga fulva Airy Shaw, Kew Bull. 19 (1965) 321; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 154.
Macaranga repando-dentata Airy Shaw, Kew Bull. 19 (1965) 322; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 155.
Treelets, 2 m, or small trees, to 8 m. Twigs solid, slender, 23 mm, terete, ageing rugose, variously strigose with long stiff spreading white hairs and variously fulvous tomentose. Stipules and bud scales subulate, to 10 mm, sinuous, erect, crowded at tip, sparsely strigose, somewhat persistent at nodes. Leaves: petioles 28 cm, slender, 1.5 mm diam., terete, indumentum as twig; blades ovate-oblong, (5)11(23) by (2)4.5(8) cm, papery, base rounded often cordulate rarely minutely peltate, usually with several small glands near petiole insertion, margin weakly sinuate and often with small distant glandular teeth, apex abruptly narrowing to a 2 cm acumen, drying warm (fulvous) brown to khaki green, strigose at least on nerves below, sometimes on lamina below and above, with sparse usually dark granular glands below. Staminate unknown. Infructescences crowded, spicate, axis 0.53 cm, terete, slender. indumentum as twig; bracts 2, apical, round or reniform, 20 by 25 mm diam., leaf-like, papery, blunt, margin entire or toothed, nerved, nerves strigose, enclosing a single dicoccous fruit. Fruit with pedicel to 10 mm, dicoccous, lobes to 8 mm diam., woody, densely fulvous tomentose with a few 1 mm apical spines; styles 2, tomentellose, caducous, thread-like, smooth, to 25 mm. Seeds globose, 5 mm, smooth, enclosed in a thin sarcotesta.
Distribution Throughout Borneo.
Habitat & Ecology In the understorey of primary lowland forest, on sandy soils, also including ultrabasics in Sabah. Locally gregarious. Up to 300 m.
Note Airy Shaw described this and M. fulva from one collection each and M. repando-dentata from four. Now that more material is to hand the characters which he noted distinguished them and those he used to key them out do not hold. I consider this to be just one somewhat variable species. I retain for it the most descriptive of his three epithets. The type and sole collection of M. fulva has exceptionally small leaves and short infructescences but other collections approach this state. M. strigosissima is easily recognised from the densely to sparsely strigose parts, especially petioles and twigs.
Macaranga subpeltata K.Schum.& Lauterb., Fl. Schutzgeb. Sόdsee (1900) 400; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 359; L.M.Perry, J. Arnold Arb. 34 (1953) 247; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 158; Gen. Macaranga Prodr. (2008) 251. Syntypes: Lauterbach 1023, Lauterbach 1261, Lauterbach 2635 (not seen; Lauterbach 635 at K), Papua New Guinea, Madang Prov., Gogol.
Small trees, to 15 m. Twigs fairly stout, 5 mm, terete, densely golden pubescent, sometimes glabrescent, with stipule scars. Stipules very broad, oblong-ovate 8 by 68 mm, apex acute, sometimes as an abrupt central tip, with a central rib, outside densely to sparsely pubescent, inside glabrous, papery, erect, caducous. Leaves: petioles 510 cm, 2 mm diam., terete, pubescent as twigs, kneed at top; blades ovate to trullate, (11)22 by (7)16 cm, thickly papery, base broadly rounded, broadly shallowly cordate to truncate and cordulate, with several small scattered glands near petiole insertion, with 3 major and several minor basal nerves, margin repand and with distant small glandular teeth, apex acute to acuminate, drying khaki above warm brown below, above densely tomentose on main nerves and lamina scabrid, below simple and tufted hairs on all nerves densest on major ones nearly always without granular glands. Staminate inflorescences coarse sometimes large panicles with 34 axis orders, 620 by 410 cm, pubescence as twigs; branches ascending; peduncle 13 cm; main axis 13 mm diam., terete; branches alternate, secondaries to 14 cm, tertiaries to 6 cm; bracts ovate, to 6 by 3 mm, apex acute, papery, slightly pubescent, persistent; flower clusters crowded in coarse loose catkins to 15 by 5 mm at branch tips; bracteoles ovate c. 5 by mm plus 1 mm wide base, apex acute, coarsely deeply distantly dentate, surfaces plus margins with sparse hairs, pinnately nerved, at margins nerves ending in a gland, papery, erect, persistent, much longer than cluster. Staminate flowers 46 per cluster; stamens 4(?), anthers 4-locular. Infructescences spikes to 10 cm or sparsely branched racemes to 26 cm with 6 cm branches and 8 cm peduncle; main axis stout somewhat flattened, 3 mm diam., pubescence as twigs; bracts at branches persistent, as staminate; fruits in clusters of 14 at branch tips, subtended by pinnately nerved 5 mm stalked leafy bracts, ovate, to 2 by 1 cm, apex acute, papery, abaxially sparsely pubescent and sparsely dark granular glandular, adaxially more or less glabrous. Fruits strongly dicoccous, 6 by 11 mm, thinly woody, densely golden pubescent and with a few dark granular glands with at least on upper part dense soft 4 mm pubescent spines tapering from a swollen base, sessile; calyx persistent; style short, plumose, persistent. Seeds round, very coarsely verrucose, with sarcotesta.
Distribution East New Guinea (all modern collections Morobe District).
Habitat & Ecology Primary of secondary forest, 300800 m.
Note Distinctive in its strongly dicoccous fruits, big leaflike dentate staminate bracteoles, and usual absence of granular glands on leaves below.
Macaranga suleensis Whitmore, Kew Bull. 34 (1979) 605; in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 158; Gen. Macaranga Prodr. (2008) 252. Type: Croft & Katik NGF 14969, (holo K; iso L), Papua New Guinea, East New Britain Distr., Pomio Subdistr., Mt. Sule, c. 25 miles NNE of Fullerborn Harbour.
Small tree, 5 m. Twigs slender, 2 mm diam., terete, striate, glabrous. Stipules elliptic, 5 by 1.5 mm, soon caducous. Leaves: petioles c. 3 cm, slender, 1 mm diam., terete, glabrous, weakly kneed at top; blades oblong-elliptic, c. 10 by 2.5 cm, papery, narrowing gradually to a rounded base with 2 small glands near petiole insertion and to a long acuminate apex, margin repand to crenate, drying grey green, glabrous, without granular glands, secondary nerves slender, 810 pairs, running straight and parallel almost to margin, domatia absent. Staminate absent. Pistillate inflorescences amongst leaves, tiny, delicate 3 cm glabrous spikes; peduncle 2 mm, axis thread-like, terete with evenly spaced distant solitary flowers; bracteoles subtending flowers, triangular, 0.8 mm, persistent, spreading, glabrous. Pistillate flowers: pedicel 13 mm; sepals 2, free, ovate, 1 mm, granular glandular; ovary golden granular-glandular, style lateral, 3 mm, recurved, papillose. Fruits and Seeds unknown.
Distribution Bismarck Archipelago (New Britain; known from type only).
Habitat & Ecology Montane forest, 1500 m.
Note Similar to M. gracilis but with quite different nervation.
Macaranga sumatrana Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1003; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 238; Whitmore in Airy Shaw, Kew Bull. 36 (1981) 322; Gen. Macaranga Prodr. (2008) 252. Type: Korthals s.n. (holo G-DC; iso L).
Macaranga necopina Whitmore, Kew Bull. 36 (1981) 423; in Airy Shaw, Kew Bull. 36 (1981) 322.
Trees, to 18 m. Twigs slender, quadrangular, 3 mm diam., blackish, rusty tomentose at tip. Stipules ovate, 6 by 4 mm, apex acute to acuminate, papery, with a central nerve, erect, subpersistent. Leaves: petioles 69 cm, terete, 1.5 mm diam., finely golden pubescent, slightly kneed at top; blades deltoid, to c. 16 by 6 cm, thinly leathery, base broadly truncate, not peltate, 23 small glands at petiole insertion, margin sinuate with very sparse emergent glandular teeth, apex broadly acute, drying khaki with nerves brown, on both surfaces secondaries and tertiaries (and sometimes lamina also on lower surface) with short close simple and tufted hairs, lower surface with close black granular glands, secondary nerves straight, parallel, basal pair at petiole insertion, tertiary nerves scalariform. Staminate inflorescences amongst and behind leaves, narrow crowded panicles, c. 13cm long, with 3 tawny pubescent axis orders; secondary axes divaricate, slightly ascending, 23 cm long; peduncle 4 cm long, fairly slender, 1.5 mm diam., terete, ridged; secondaries with bracts; latter trapezoid, 4 by 2.5 mm, broadest near middle, broadly acute, irregularly coarsely deeply dentate, pubescent, subtending a 5 mm axis of crowded overlapping bracteoles; latter broadly ovate, reaching 2 by 3 mm, long narrow 1 mm glandular tip, margins with regular blunt teeth, each tipped by a small gland, concave, pubescent both sides, thick texture, erect, persistent, longer than cluster. Staminate flowers c. 10 per cluster, tiny round, 0.2 mm diam.; pedicel slender, 0.3 mm; sepals 2, pubescent; stamens 2, anthers 4-locular. Pistillate inflorescences behind leaves, narrow, 15 by 6 cm, rusty pubescent racemes; peduncle 5 cm, terete; branches 3 cm, normal to axis. Fruits in clusters of 35, small, weakly bilocular, 2 by 3 mm, thinly woody, densely black granular glandular; pexicel slender, 4 mm; calyx small, persistent; styles 2, 1 mm, caducous, spreading, apical, fused at base. Seeds globose, 1.5 mm diam., with sarcotesta.
Distribution Sumatra (Aceh, Jambi, Indrapura), endemic, apparently uncommon.
Habitat & Ecology Secondary lowland forests.
Macaranga suwo Whitmore, Kew Bull. 34 (1979) 605; in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 159; Gen. Macaranga Prodr. (2008) 253. Type: Schram BW 13263 (holo K; iso L), Indonesia, Papua, Vogelkop, Warsamson Valley, E of Sorong.
Buttressed tree, 10(30) m tall, clear bole 8 m, diam. 40 cm, developing red heartwood. Twigs strongly angular, 7 mm across, with conspicuous stipule scars, usually glaucous grey, sparsely hispid or glabrous. Stipules united, apparently single, as a 4(8) cm apical bud, long tapering, 9 mm diam., gibbous towards base, rusty hispid, leathery, caducous. Leaves: petioles 9 cm, flattened above near base, 3 mm diam., finely tomentose plus sparse long hairs, rarely glabrous, slightly glaucous, kneed at both ends; blades big, ovate to ovate-elliptic, c. 26 by 15 cm, leathery, base broadly rounded to subtruncate 1 cm peltate, margin repand, apex acuminate, sometimes tricusped or with small lateral lobes, usually glaucous grey below with the nerves pale brown, above glabrous, shiny, below finely velvety plus a few long hairs, nerves very close and wholly covered in tiny clustered hairs, granular glands patchy, lamina-coloured, almost invisible; secondary nerves pinnate, basal pair from petiole insertion and with strong scalariform tertiaries. Staminate inflorescences broad racemes, 7 by 6 cm; main axis terete or slightly angular, 1 mm diam., tawny hispid; peduncle 6 mm; branches spreading, 6 mm apart, to 4 cm long, sinuous, ends thread-like; flower clusters crowded at tips or regularly 3 mm spaced; bracts stipule-like, elliptic, to 10 by 3 mm, outside rusty hispid, caducous; flower clusters crowded; bracteoles triangular, 3 by 3 mm, apex acute, strongly cucullate, entire, pubescent and with a few long hairs without, glabrous within, thin, enclosing cluster, erect, persistent. Staminate flowers c. 12 per cluster, minute, shortly pedicellate; stamens 24, anthers 4-locular. Infructescences 722 cm racemes; axis stout, strongly flattened, to 3 mm across, coarsely striate, slightly glaucous, densely to sparsely rusty hispid; peduncle 3.512 cm; sparse branches to 15 mm, adpressed to main axis, alternate or opposite; bracts stipule-like, triangular, 20 mm, regularly tapering from broad 7 mm wide base, leathery, slightly glaucous, sparsely hispid, caducous, leaving a scar. Fruits sessile, at first wholly enclosed in 3 mm diam. globular glaucous calyx with 2 recurved densely papillose 4 mm styles protruding; at maturity depressed globose, 3 cm diam., surface like a hedgehog, densely covered with glaucous grey pubescent broad tapering sinuous 12 mm long spines, circumscissile, the top half falling to expose 3 loculi set in 2 mm thick woody pericarp tissue; calyx persistent. Seeds globose, 3 mm diam., coarsely verrucose, with sarcotesta.
Distribution West New Guinea (Sorong, common in Ayawasi).
Habitat & Ecology Secondary forest, sometimes on white sand, 50460 m.
Notes 1. This becomes a fair sized forest tree with hard wood but has only a short clear bole.
2. Reminiscent of M. hispida in the twigs and clustered leaf hairs. Unusual in this species group in not having patellar glands on the bracteoles. The circumscissile capsules are bizarre and unique.
Macaranga sylvatica Elmer, Leafl. Philipp. Bot. 2 (1908) 431; Merr., Enum. Philipp. Flow. Pl. (1923) 443; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 35; Whitmore, Gen. Macaranga Prodr. (2008) 254. Macaranga ramiflora Elmer var. sylvatica (Elmer) Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 324. Syntypes: Elmer 9824 (BO, K, L, W), Elmer 9838 (BO, K, L, W), Elmer 10361 (BO, W), all Philippines, Negros, Cuernos Mountains, Dumguete.
Macaranga ramiflora Elmer, Leafl. Philipp. Bot. 2 (1908) 427; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 35.
Macaranga cuernosensis Elmer, Leafl. Philipp. Bot. 2 (1908) 429; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 34.
Macaranga montana Merr., Philipp. J. Sc. Bot. 7 (1912) 394 [non Heyne ex Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii, 1914]; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 35.
Macaranga merrilliana Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 334.
Trees, to 1015 m tall. Twigs with conspicuous leaf scars, usually slightly angular, furfuraceous. Stipules elliptic, 69 mm long, usually finely furfuraceous, papery, chocolate colour, usually soon caducous. Leaves: petioles (2)59 cm, slender, rounded, striate, glabrescent; blades ovate or occasionally elliptic, 616 by 210 cm, papery, base cuneate to truncate, sometimes with small glands near petiole insertion, apex acuminate, below with tiny usually sparse granular glands and often nerves pubescent with dense fine white hairs, secondary nerves c.12 pairs. Inflorescences crowded, often dense, tangled, short, to 6 cm, axes thread-like, scurfy, not or sparsely once shortly branched, with grouped or evenly spaced and close small 23 mm flower clusters; bracteoles tiny, under 12 mm, apex acute, furfuraceous, glandless, persistent. Staminate flowers (2)6(8) per cluster; sepals free; stamens (4)6(13), anthers 4-locular. Fruits solitary or up to 4, round or slightly bilobed, 3 mm, thinly woody, smooth, slightly furfuraceous; pedicel 25 mm long, thread-like; calyx tiny, persistent; styles 2, barbed, thread-like, apical or slightly off-centre, as long as fruit, soon caducous. Seeds round, verrucose.
Distribution Philippines (Luzon, Samar, Mindoro, Leyte, Negros, Mindanao).
Habitat & Ecology Mountains, (500)10001200(2200) m.
Note The distinctive characters are the tiny persistent eglandular bracteoles, the fruits often solitary with slender (caducous) styles as long as their diameter. This is a montane species. Somewhat variable, including in the characters used to distinguish the several species that I now reduce. As with M. cumingii (see there) careful field study might just possibly show that more than one discrete taxon exists. The type of M. montana (Merrill 8076) does not have a lateral style (Merrill 1912), however PPI 3229 does have the usually caducous style slightly eccentric, otherwise it matches this species.
Macaranga tanarius (L.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 997; Benth., Fl. Austr. 6 (1873) 146; F.M.Bailey, Fl. Queensland 5 (1902) 1452; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 352, t. 59; Pflanzenr. IV.147.xiv add. vi (1919) 28; Pflanzenr. IV.147.xvii add. vii (1924) 184; Merr., Interp. Herb. Amboin. (1917) 320; Enum. Philipp. Flow. Pl. (1923) 443; Ridl., Fl. Malay. Penins. 3 (1924); Backer & Bakh.f., Fl. Java 1 (1963) 488; Airy Shaw, Kew Bull. 23 (1969) 99; Kew Bull. 26 (1971) 291; Kew Bull. 31 (1976) 395; Whitmore, Gard. Bull. Singapore 31 (1978) 56; Tree Fl. Malaya 2 (1973) 111; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 158; in Airy Shaw, Kew Bull. 36 (1981) 322; Chiu, Guihaia 2 (1982)147, in clavi; Airy Shaw, Alph. Enum. Philipp. Is. (1983) 35; Chakrab. & N.P.Balakr., J. Econ. Taxon. Bot. Addit. Ser. 9(1992) 81; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 379, Fig. 19, Plate XX: 2; Whitmore, Gen. Macaranga Prodr. (2008) 255. Tanarius minor alba Rumph., Herb. Ambon. 3 (17411745) 190, t. 121, nom. inval. Ricinus tanarius L. in Stickm., Herb. Amboin. (1754) 14; Amoen. Acad. 4 (1759) 125. Mappa tanarius (L.) Spreng., Syst. Veg. 3 (1826) 878. Macaranga tanarius var. genuina Mόll.Arg. in DC., Prodr. 15, 2 (1866), 997, nom. inval. Type: Rumphius (17411745) 190, t. 121 (see note 2).
Tanarius minor rubra Rumph., Herb. Ambon. 3 (17411745) 190, nom. inval.
Ricinus mappa Roxb., Fl. Ind. ed. 2 (1832) 690, non L. (see Merrill, 1917).
Mappa glabra Juss., Nouv. Ann. Hist. Nat. 3 (1834) 487; Euphorb. Gen. Tent. (1824) 114, Pl. 14, fig. 44; Merr., Interp. Herb. Amboin. (1917) 321 passim; Airy Shaw, Kew Bull. 37 (1982) 26. Macaranga glabra (Juss.) Pax & Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 355.
[Macaranga vulcanica Elmer ex Merr., Enum. Philipp. Flow. Pl. (1923) 443, nom. nud.].
var. tanarius
var. tanarius: Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 380; Whitmore, Gen. Macaranga Prodr. (2008) 256.
Small trees. Twigs terete, glabrous. Stipules ovate, to 15 by 8 mm, apex acute, glabrous, dark brown, papery, erect, subpersistent or caducous. Expanding leaves: petioles and leaves pubescent, very soon becoming glabrous except at base of main nerves below. Leaves: petioles to 15 cm, terete; blades ovate, to 17 by 15 cm, papery, base rounded, to 4 cm peltate, margin distantly finely toothed, apex acute to acuminate, usually drying khaki brown, glabrous, below with usually dark, rarely golden, granular glands, with spider's web nervation. Staminate inflorescences variable; narrow panicles to 15 cm long, glabrous, main axis flattened; basal peduncle to 10 cm; secondary axes in fascicles of 35, lowest usually subopposite, tertiary thread-like axes; flower clusters spaced; branches subtended by usually soon caducous stipule-like bracts, to 8 mm long; bracteoles persistent, larger than clusters, to 6 by 3 mm, apex acuminate to caudate, margin toothed, glabrous, sometimes nerved; clusters developing more or less simultaneously. Staminate flowers 56(14) per cluster, tiny, 0.5 mm; pedicel 11.5 mm; sepals free, glabrous; stamens 67, anthers 4-locular. Infructescences with fruits crowded at head of a 416 cm, bare, flattened, glabrous peduncle, subtended by caducous, sometimes stalked bracts; latter leafy, ovate, 4 by 4 mm or larger, apex acuminate, upper margin deeply dentate to fimbriate, papery, glabrous, concave. Fruits bilobed, to 10 mm across by 6 mm, leathery, glabrous, with numerous long slender processes from upper half, woody; pedicel 2 mm; calyx tiny, persistent; styles 2, persistent, 5 mm, filiform, apical. Seeds round, verrucose.
Distribution Thailand (rare), S. Sumatra, Java, Borneo, Philippines (poorly collected but the commonest variety, Merrill 1923), Sulawesi, Lesser Sunda Islands (Timor), Moluccas (Ternate, Ceram, Ambon), New Guinea, Australia (Queensland), Solomon Islands (Guadalcanal, Malaita), Vanuatu (widespread).
Note Much less common than var. tomentosa.
var. tomentosa (Blume) Mόll.Arg.
var. tomentosa (Blume) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 997; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 3380, Fig. 19; Whitmore, Gen. Macaranga Prodr. (2008) 257, Fig. 22. Mappa tomentosa Blume, Bijdr. (1825) 624; Baill., Ιtude Euphorb. (1858) 429; Atlas t. 20, f.14. Rottlera tomentosa (Blume) Hassk., Cat. Hort. Bogor (1844) 238. Type: Cuming 684 (iso: CGE, G, OXF, K, W) [more or less lectotypified by Whitmore 2008].
Croton lacciferum Blanco, Fl. Filip. (1837) 752; Fl. Filip., ed. 2 (1845) 517; Fl. Filip., ed. 3, 3 (1879) 153; Merr., Sp. Blancoan. (1918) 226.
Macaranga tanarius (Blume) Mόll.Arg. var. brevibracteata Mόll.Arg. in DC., Prodr. 15, 2 (1866) 998.
Macaranga molliuscula Kurz, J. Asiat. Soc. Bengal 42 (1873) 245.
As var. tanarius except: Twigs finely pubescent or furfuraceous (sometimes becoming glabrous). Stipules broadly elliptic, 820 by 37 mm, apex acuminate to caudate, puberulous or furfuraceous, sometimes just on midline, sometimes becoming glabrous. Leaves: petioles slightly furfuraceous or pubescent, at least at tip; blades to 17 by 15 cm, usually smaller, drying dark chocolate brown, furfurescent to pubescent on nerves below, sometimes just near petiole insertion, granular glands minute, inconspicuous, sparse to dense, golden or blackish. Staminate inflorescences 2 to 3 orders of furfuraceous axes, sometimes with axillary branches; bracteoles ovate, size very variable, 1.5(12) by 1.5(13) mm, apex acute to acuminate or caudate, margin usually distantly deeply toothed, both surfaces finely pubescent or rarely (almost) glabrous, concave, papery, with a conspicuous network of nerves, usually overlapping, subtending clusters of 56(14) flowers developing in sequence, and from which a next order axis often later developing; clusters grouped. Staminate flowers: sepals pubescent. Infructescences with peduncle and bracts furfuraceous. Fruits variously granular glandular.
Distribution India (including Andaman and Nicobar Islands), Burma (Tenasserim), China, Taiwan, Japan (Ryukyu Islands), Thailand, Malay Peninsula, Sumatra (incl. Anambas Islands), Java, Christmas Island (Indian Ocean), Borneo (absent around Kuching in Sarawak), Philippines (much collected, but the less common variety; Merrill 1923), Sulawesi, Lesser Sunda Islands (Bali, Timor, West Sumbawa, Sumba, Wetar), Moluccas (Tanimbar, Aru, Seram), New Guinea, Australia (Northern Territory, Queensland, New South Wales), Solomon Islands, Vanuata, Kiribati (Tarawa: cult.).
Habitat & Ecology Very common. Slik et al.: Usually in very disturbed vegetation like scrub or abandoned shifting cultivation areas, often along roads, or along streams. On clay loam, sandy, and sandstone soils. Up to 2100 m altitude.
Notes 1. Macaranga tanarius is the most widespread of all species in the genus, ranging from India to China and Ryukyu, south through Thailand, and throughout Malesia to New South Wales (but which variety?), the Solomons and Vanuatu. It is polymorphic. There is a completely glabrous form (though this has the expanding petioles and leaves pubescent) and variously tomentose forms which collectively are much commoner.
2. Merrill in his interpretation of Rumphius' Herbarium Amboinensis (1917: 320321) was in no doubt that the glabrous form matches the Rumphius plate, which is the type of Ricinus tanarius L. and was of the view that the taxon is quite distinct from the pubescent form which he knew well in the Philippines. This glabrous taxon therefore becomes the type variety of the species. Because the two forms approach so closely, I do not consider the much commoner tomentose one can be given more than varietal status and I do not share the view of Pax & Hoffman (1914: 354) that no varieties can be distinguished. The tomentose form is itself highly variable but I have been unable to distinguish discrete entities within it. Apart from variation in degree of hairiness the staminate bracteoles vary substantially in size, from 1.5 by 1.5 mm to 12 by 13 mm, sometimes within a single locality, for which reason I do not recognise var. brevibracteata of Mόll.Arg. (which is the commonest form in the Solomon Islands and Vanuatu). The pistillate bracteoles are never so small as they are in some staminate specimens.
3. Pax & Hoffmann (1914: 354) stated: A species with very variable leaf indumentum but which scarcely may be divided into truly limited varieties. Smith [1910: 409] says that Mόllers varieties (α tomentosum, β genuinum) are scarcely distinct. All authors describe leaves as eglandules above, nevertheless well-developed maculate glands have been observed in specimens (Java, Koorders 14084β; New Guinea, Weinland 8100; Philippines, Fιnix 476, 3990, Jager et al. s.n.). Such glands are not of constant appearance. Male bracts sometimes bigger and smaller; f. brevibracteata Mόll.Arg. is the condition with small bracts, joined to all the others.
Macaranga tentaculata Airy Shaw, Kew Bull. 23 (1969) 102; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 159; Gen. Macaranga Prodr. (2008) 259. Type: Brass 24988 (holo K; iso L), Papua New Guinea, Milne Bay Distr., DEntrecasteaux Islands, Goodenough Island.
Small tree to 12 m. Twigs stout, to 10 mm diam., angled, tomentose. Stipules long triangular, to 25 by 9 mm, papery, with straight sides, tapering from base to acuminate tip, glabrous except for a central nerve, sparsely hairy on its abaxial surface, mid or dark brown, recurved, persistent. Leaves: petioles to 21 cm, 5 mm diam., terete, finely striate, tomentose, drying with basal 1 cm strongly constricted; blades oblong-ovate, to 30 by 23 cm, papery, base broadly rounded deeply 7 cm peltate, margin repand, apex abruptly 23 cm caudate, drying mid brown, below velvety (densely so when young), with dense spreading white hairs on all nerves, and somewhat patchy tiny golden granular glands, above with sparse long white hairs, spider's web nervation with scalariform tertiaries. Staminate unknown. Infructescences small, axillary 8 cm racemes, main axis stout somewhat flattened, 3 mm diam., tomentose; peduncle 3 cm. Fruits solitary, crowded towards branch tips, bilobed, 6 by 11 mm, with sparse reddish granular glands, densely beset with slender sinuous reddish 68 mm processes bearing a small swollen apical gland, thinly woody; pedicel stout, 5 mm; calyx persistent; styles 2, subulate, 56 mm, papillose. Seeds (immature) ellipsoid, 6 by 4 mm, coarsely shallowly verrucose, with sarcotesta.
Distribution East New Guinea (Goodenough Island), Bismarck Archipelago (W. New Britain).
Habitat & Ecology Lowlands at 150800 m.
Notes 1. The New Britain collection (Ridsdale NGF 30425) differs from the type in having the fruit tentacles mostly glandless.
2. Somewhat similar in fruit to M. clavata and M. herculis (see there).
Macaranga tessellata Gage, Nova Guinea Bot. 12 (1917) 481, t. 186; Pax & K.Hoffm. in Engl.,Pflanzenr. IV.147.xvii add. vii (1924) 184; L.M.Perry, J. Arnold Arb. 34 (1953) 207; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 159; Gen. Macaranga Prodr. (2008) 260. Syntypes: Pulle 132 (BO, L, K); Pulle 177 (L), both Indonesia, Papua, Noordfluss, near Kloofbivak.
Trees, to 16 m. Twigs slender, 4 mm diam., terete, with conspicuous stipule scars, densely yellowish brown pubescent. Stipules united, apprently single as a sinuous elliptic apical bud, 39 by 0.51.2 cm, acute, papery, dull yellowish brown sericeous, erect soon caducous. Leaves: petioles 38 cm, slender, 12 mm diam., terete, pubescent, kneed at top; blades ovate, 1528 by 813 cm, papery, base broadly rounded, cordulate, and sometimes with glands at petiole insertion, margin repand, apex an abrupt 1 cm caudex, drying mid grey brown above, slightly golden brown below, above glabrous except pubescent along midrib near base, below densely pubescent on nerves and with well spaced black granular glands, pinnately nerved with tertiaries and quaternaries scalariform. Staminate inflorescences delicate sparsely branched 711 cm densely pubescent racemes; main axis flattened, 1.5 mm across; peduncle 27 cm; branches sometimes opposite, sometimes with short axillary branches, bearing up to 5 mm spaced tiny 1 mm diam. flower clusters, sometimes zigzag; bracteoles 2 mm long, spreading, longer than cluster, with a 1 mm broad base narrowing abruptly into an oblong abruptly acuminate limb bearing adaxially numerous patellar glands, and pubescent abaxially, somewhat fleshy. Staminate flowers c. 8 per cluster, tiny, subsessile; stamens 3, anthers 3-locular. Infructescences spicate, 7 cm long; axis stout, 2 mm across, slightly flattened, densely pubescent, bearing a single apical weakly bilobed fruit. Fruits 10 by 17 mm, densely covered in 3 mm soft golden sericeous flattish spines, sessile; calyx caducous. Seeds globose, 5 mm, irregularly warty.
Distribution New Guinea.
Habitat & Ecology Primary forest, 30800 m.
Macaranga thomasii Whitmore, Gen. Macaranga Prodr. (2008) 260. Type: Kostermans 1224 (holo K; iso L, P), Moluccas, Morotai, Gunung Pare Pare.
Small trees or to 30 m. Twigs (fairly) slender, 2(5) mm diam., strongly angular, somewhat flattened towards tip, drying blackish brown, glabrous, with conspicuous stipule scars. Stipules unites, apparently single, forming a slightly sinuous elliptic apical bud 45 cm long, lanate with long adpressed grey brown hairs, papery. Leaves: petioles 413 cm, slender, angular, glabrous, blackish brown; blades ovate-oblong, 1235 by 520 cm, thinly leathery, base rounded, cordulate, with several glands, margin shallowly sinuate, revolute, apex acuminate, drying grey green above pale brown below, finely pubescent at least on main nerves below and with fairly distant dark granular glands, secondary nerves 1012 pairs. Staminate unknown. Infructescences spikes, to 12 cm, with a single terminal fruit, axis slender, flattened, globose; bracteoles soon caducous, elliptic, to 9 by 1 mm including 4 mm stalk, at first scurfy, with small patellar glands, margin sinuous, apex tipped by gland. Fruits depressed globose, 10 by 8 mm, weakly bilobed, covered in flattened slightly hooked slender spines, surface finely granular glandular; pedicel 510 mm; calyx at first enclosing fruit in papery evanescent golden tomentose lobes; style soon caducous, 4 mm, bearing 2 linear, 6 mm, erect, papillose stigmas. Seeds round, 6 mm diam. , coarsely irregularly prominently verrucose, with thin sarcotesta.
Distribution Moluccas (Halmahera, Morotai).
Habitat & Ecology Locally abundant, 6001000 m.
Notes 1. Similar to M. longistipulata, differing principally in the longer petiole, fewer secondary nerves, glands at leaf base and longer fruiting spike.
2. This species is named for my son Tom in whose company it was collected.
Macaranga trachyphylla Airy Shaw, Kew Bull. 25 (1971) 534; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 150; S.J.Davies, Harvard Pap. Bot. 6 (2001) 417. Type: Richards 1437 (holotype: K), Malaysia, Sarawak, Gunung Dulit.
Subcanopy pioneer tree to 20 m tall with a candelabra-shaped crown; twigs 712 mm in diam., evenly pubescent with fine silvery hooked coarse hairs, hollow, housing ants. Bark smooth, grey-brown, hoop-marked. Stipules ovate, 79 by 1014 mm, recurved, succulent, dark purplish brown to almost black, producing food-bodies on the abaxial surface, 36 pairs present on shoots. Leaves: petioles terete, stout, 1840 cm long, finely pubescent with hooked coarse hairs; blades broadly to very broadly ovate, large to very large and variable in size, 2545 by 2040 cm, trilobed, dissected to 1/31/2 of the leaf length, lateral lobes broad, sinuses narrow, >4 cm peltate, base very broadly rounded, margin entire, rarely with small protruding nectaries, apices broadly acute, evenly pubescent on both surfaces, adaxial surface rough to touch due to the coarse hooked hairs, not glaucous; 1° venation palmate with 78 prominent veins, 2°4° venation scalariform, 2° veins curved upwards just before the leaf margin, minor veins densely reticulate; young leaves red-brown, pubescent. Staminate inflorescences paniculate, erect, 1521 by 613 cm, covered in coarse hooked silvery hairs, to 4 axis orders, main axis flattened, first pair of secondary branches ± opposite with accessory branches; bracts ovate-elliptic, 1014 by 35 mm, margin entire, apex acute, very finely tomentose, caducous; flower clusters with c. 20 flowers, spirally arranged and grouped towards the distal end of ultimate inflorescence branches; bracteoles ovate-elliptic, 68 by 23 mm, enclosing flower clusters, margin entire, apex caudate, glabrous or rarely with scattered minute red-brown hairs. Staminate flowers c. 1 mm long, sessile; sepals fused, splitting irregularly to c. 1/3, apex densely covered in minute red-brown hairs; stamens 1; anthers 3-locular. Pistillate inflorescences paniculate, erect, 815 by 513 cm, stout, with coarse hooked silvery hairs, up to 3 axis orders; bracts caducous, not seen. Pistillate flowers c. 56 mm long, solitary in bracteole axils; calyx urceolate, c. 23 mm long, apex truncate with tiny silvery hooked hairs, persistent, splitting irregularly as ovary expands; ovary (4)5-carpellate, 23 mm long; styles 23 mm long, fused for 3/4 of length, free at the apex, persisting to form a prominent 34 mm long crown at the fruit apex; stigma not dissected. Fruits subglobose, 56 by 911 mm, sessile, one discrete glandular patch on each carpel wall developing into a slender horn-like process 23 mm long covered in yellowish-green sticky exudate. Seeds subtriangular-ovoid, 3.54 mm in diam., black, shallowly pitted and grooved, with a small cruciform scar, encased in a fleshy pale reddish-purple aril.
Distribution Borneo (Brunei, Sabah and Sarawak). It has been collected from Gunung Gading in the far west of Sarawak throughout Sarawak and Brunei to Penampang in the south western corner of Sabah.
Habitat & Ecology Macaranga trachyphylla is a common early successional tree in both primary and secondary mixed dipterocarp forest in the lowlands of west Borneo up to c. 500 m. It is moderately shade tolerant and is common in canopy openings along streams in primary forest. It is typically found on moderately nutrient-rich soils as in Bukit Belalong in Brunei and on the shale-derived soils at Lambir Hills National Park. It is mostly absent from swampy or flooded areas and very degraded successional sites.
Notes 1. The rough adaxial leaf surfaces that are "almost as rough as a piece of sandpaper" (Airy Shaw 1971) are diagnostic for this species. No other member of section Pachystemon has such rough leaf surfaces. The stout twigs have a similarly rough indumentum. The elegant candelabra crowns of M. trachyphylla are at once recognizable with the large trilobed pendant leaves. The stipules, whilst similar to several other myrmecophytic members of section Pachystemon, are distinctive in typically being dark purple to almost black when fresh. A further distinctive feature is the caudate eglandular apices of the staminate bracteoles, which is not found in closely related species.
2. It is probably most closely related to M. indistincta from eastern Borneo, with which it shares a similar ecology and fruit morphology. However, rough leaf surfaces and caudate staminate bracteoles are not found in M. indistincta.
Macaranga trichanthera L.M.Perry, J. Arnold Arb. 34 (1953) 243; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 160; Gen. Macaranga Prodr. (2008) 262. Type: Brass 5077 (holo A; iso K), Papua New Guinea, Central Prov., Mt. Tafa.
Trees, 625 m. Twigs 3 mm diam., angled, densely finely rusty brown pubescent. Stipules elliptic, 6 by 2 mm, apex acute, papery, erect, densely pubescent, soon caducous. Leaves: petioles 24 cm, diam. 2 mm, terete, striate, finely rusty pubescent becoming glabrous, kneed at both ends; blades ovate, 813.5 by 510 cm, leathery, base broadly rounded, 110 mm peltate, basal secondary nerves and some minor nerves from petiole insertion, margin slightly repand, apex with an abrupt 1 cm acumen, drying grey above, becoming glabrous, below densely rusty brown tomentum obscuring black granular glands, secondaries and reticulations very slightly raised above, strongly raised below, finely bullate. Staminate inflorescences small 4 cm sparsely branched rusty pubescent racemes; branches to 15 mm, alternate, divaricate, arising from a flower cluster; peduncle 510 mm; main axis terete, 1.5 mm diam.; flower clusters mostly contiguous; bracteoles tiny, obovate, 1 by 0.8 mm, apex blunt, glabrous, smaller than cluster, persistent. Staminate flowers in clusters of 15; calyx hairy; stamens 35, anthers 4-locular. Pistillate inflorescences spikes to 8 cm. Pistillate flowers sessile; calyx 2-lobed; ovary 2-locular, pubescent and granular glandular; styles 2, joined at base, 6 mm long, papillose. Fruits globose, 1 cm, leathery. Seeds rugulose.
Distribution Throughout New Guinea.
Habitat & Ecology Primary or less often secondary montane forest, 18002400 m.
Notes 1. Pistillate not seen, description from Perry (1953), pace whom no hairs were seen on the anthers.
2. Leaves differ from M. albescens in absence of a basal lobe.
Macaranga trichocarpa (Zoll.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1003; Hook.f., Fl. Brit. India 5 (1887) 450; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 358; Ridl., Fl. Malay. Penins. 3 (1924) 303; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 441; Corner, Ways. Trees Malaya (1940) 268; Whitmore, Tree Fl. Malaya 2 (1973) 111; in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 158; Gard. Bull. Singapore 31 (1978) 56; in Airy Shaw, Kew Bull. 36 (1981) 322; Airy Shaw, Kew Bull. 23 (1969) 97; Kew Bull. 26 (1972) 291; Chakrab. & N.P.Balakr., J. Econ. Taxon. Bot. Addit. Ser. 9 (1992) 82; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 382, Fig. 20; Whitmore, Gen. Macaranga Prodr. (2008) 263; Fig. 23. Mappa trichocarpa Zoll., Linnaea 28 (1856) 307; Verhand. Natuurk. Vereen. Ned.-Indiλ 1 (1856) 8. Type: Zollinger & Moritzi 3045 (not seen), Sumatra, near Lampong.
Mappa zollingeriana Miq., Fl. Ind. Bat. Suppl. (1861) 457.
Macaranga minutiflora Mόll.Arg., Flora 47 (1864) 466; in DC., Prodr. 15, 2 (1866) 1012; Kurz, For. Fl. Brit.Burma 2 (1877) 388; Brandis, Indian Trees (1906) 592.
Mappa borneensis Mόll.Arg., Flora 47 (1864) 468, Macaranga borneensis (Mόll.Arg.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1003.
Macaranga helferi Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1004.
Macaranga trichocarpa (Zoll.) Mόll.Arg. var. trilobulata Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 442.
Bushy trees, to 6 m, with sprawling limbs (sarmentose). Bark smooth, orange brown, latex clear turning red. Twigs solid, slender, 3 mm diam., terete, smooth, finely pubescent. Stipules elliptic, 5 mm, papery, pubescent, erect, very soon caducous. Leaves: petioles 45 cm, slender, terete, finely pubescent, sometimes weakly kneed at top; blades rhombic-ovate, very occasionally with 2 lateral cusps or small lobes, 714(17) by 410(15) cm, thinly leathery, base truncate to very broadly rounded, often under 1 mm peltate, occasionally with inconspicuous glands on nerves near petiole insertion, margin with small glandular teeth, apex acute or acuminate, drying khaki below and often dark brown above, below granular glandular and velvety with short spreading white hairs on all nerves including the raised reticulum, with 3 main and several minor nerves at petiole insertion, surface above with white hairs (caducous except on nerves) arising from small persistent pustules. Inflorescences diffuse narrow racemes to 6 cm; branches sparse to 15 mm; peduncle short, 1 cm; axes slender, main axis terete, finely pubescent, lower branches alternate; bracts elliptic, 5 mm, dentate, leaf-like, pubescent, with pinnate nervation, sometimes stalked, soon caducous; flower clusters in groups, smaller than bracteoles, overlapping, branches developing through flower clusters (as M. tanarius); bracteoles persistent, ovate, strongly concave, to 4 by 2mm, apex acute, pectinate, pubescent on both sides, nerved. Staminate flowers c. 14 per cluster, maturing in sequence; pedicel 0.5 mm, tomentose; sepals united, elliptic, tomentose; stamens 2, strongly exserted, 0.5 mm long, anthers 4-locular. Pistillate flowers similar [to staminate?], ovary bilobed. Fruits round (or bilobed), 4 mm diam., woody, golden granular glandular, with close short spines bearing conspicuous glassy irritant hairs; pedicel 3 mm, pubescent; calyx small, persistent. Seeds round, 2 mm, very finely verrucose, with sarcotesta.
Distribution Lower Burma, India (Andaman Islands), Thailand (Chanthaburi only), Vietnam (Cochinchine, Annam), Malay Peninsula, Sumatra, Borneo.
Habitat & Ecology Sometimes forming thickets. In continental Asia very patchy. Usually in very disturbed secondary forest, especially scrub or abandoned shifting cultivation areas, but also present in gaps in primary forest. Often along forest edges and road sides where it can form pure stands. On sandy loam, sandy clay and red-yellow podsolic soils. Up to 500 m altitude.
Note Distinctive in habit, appearance and especially in fruit. In inflorescence construction and bracteole nervation reminiscent of M. tanarius. There is a superficial resemblance to M. kurzii.
Macaranga triloba (Thunb.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 989; Blume, Cat. (1823) 108; Hook.f., Fl. Brit. India 5 (1887) 452; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 380; Merr., Enum. Philipp. Flow. Pl. (1923) 443; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 439; Backer & Bakh.f., Fl. Java 1 (1963) 488; Airy Shaw, Kew Bull. 26 (1972) 292; Thoth. & Banerjee, Bull. Bot. Surv. India 15 (1973) 166; Chakrab., J. Econ. Taxon. Bot. Addit. Ser. 11 (1987) 22. 1987; S.J.Davies, Harvard Pap. Bot. 6 (2001) 418; Whitmore & S.J.Davies in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 384. Ricinus trilobus Thunb., Ricin. (1815) 6. Pachystemon trilobum (Thunb.) Reinw. ex Blume, Bijdr. (1825) 626. Lectotype (designated by S.J.Davies, 2001): Blume s.n. (holo: L, not seen; iso: BO, K, A [pict.]), Indonesia, Java.
Macaranga cornuta Mόll.Arg. in DC., Prodr. 15, 2 (1866) 988. Type: Teijsmann s.n. (holotype: K), Indonesia, Java, cultivated in Bogor.
Macaranga quadricornis Ridl., Kew Bull. (1923) 367. Type: Ridley s.n. (holotype: K; A [pict.]), Malaysia, Selangor, Semangkok Pass, Bukit Telaga.
Macaranga triloba auct. non (Thunb.) Mόll.Arg.L Holthuis & Lam, Blumea 5 (1942) 202; Airy Shaw, Kew Bull. 37 (1982) 28.
Small tree to 20 m tall, but frequently reproductive from as small as 34 m tall; twigs 612 mm in diam., mostly glabrous or sometimes covered in a fine tomentum of short erect hairs and then soon becoming almost glabrous, solid, not housing ants. Bark smooth, light grey-brown, hoop marked. Stipules ovate to ovate-elliptic, 715 by 514 mm, light green when fresh, spreading or slightly recurved but not appressed to the stem, coriaceous, mostly glabrous or with scattered fine ferrugineus hairs and erect silvery hairs towards the base, the pair completely encircling the twig, usually at least 4 pairs and up to 10 pairs present on the shoots. Leaves: petioles terete, (8)1025 cm long, not glaucous, glabrous or covered with short erect silvery hairs or scattered with longer erect hairs; blades ovate to broadly ovate, (12)1535 by (8)1224(30) cm, all green or with deep magenta abaxial surfaces which sometimes persist in mature leaves, trilobed, sometimes with an additional pair of basal leaf cusps, dissected to 1/32/3 of the leaf length, central lobe broad 612(14) cm wide at the base and narrowing gradually to the apex, lateral lobes usually quite broad 36(8) cm wide and ascending or spreading, 1.56.5 cm peltate, base rounded to rarely almost truncate when additional basal leaf cusps present, margin characteristically appearing almost serrate due to prominent marginal nectaries arising from 2° and 3° veins, often with scattered fine erect silvery hairs, apices acute to finely acuminate, adaxial surface mostly glabrous but often with scattered silvery erect pointed hairs at the base of the main veins, abaxial surface scattered to densely covered with short and long erect silvery hairs particularly on the veins, not glaucous; 1° venation palmate with (7)89(10) prominent veins, 2° venation scalariforin, looping near the leaf margins and terminating at the margin in prominent c. 1 mm diameter conical nectaries; 3°4° venation densely scalariform, prominent; young leaves red-brown, scattered with ferrugineus hairs and erect silvery hairs, the leaf bud very densely covered in erect sharp-pointed silvery or golden hairs which become scattered as the leaf matures, and with very prominent conical marginal nectaries. Staminate inflorescences paniculate, erect, (8)1022 by 816 cm, light green when fresh, drying black, basal axes glabrous or sometimes with scattered long silvery and short ferrugineus hairs, distal axes densely to very densely ferrugineus pubescent, 34 axis orders with the branches usually long and straight, basal unbranched main axis (1)37 cm long, flattened, first pair of secondary branches opposite with accessory branches; bracts ovate-elliptic, 58 by 36 mm, margin entire or sometimes with additional short narrow lateral cusps, apex acute to acuminate, glabrous or usually with fine ferrugineus hairs along the margin and scattered towards the base, caducous or sometimes several persisting; flower clusters with (15)2030(35) flowers, spirally arranged and grouped towards the distal ends of ultimate inflorescence branches; bracteoles ovate to ovate-elliptic, (2)35 by (2)34.5 mm, enclosing flower clusters, margin entire but sometimes dissected with 14 short lateral points, apex acuminate to finely caudate with the tip sometimes curled back, rarely acute when young, both surfaces densely covered with minute ferrugineus hairs, with a dense patch of hairs towards the base of the adaxial surface, abaxial surface usually with minute yellow granular glands. Staminate flowers 0.71.0 mm long, sessile; sepals fused, splitting into 3 lobes to 1/51/4 of length, apex densely covered in minute ferrugineus hairs; stamens 1; anthers 3-locular. Pistillate inflorescences paniculate, erect, 415 by 210 cm, stout, either unbranched or with one pair of short opposite branches, without accessory branches, the flowers usually clustered at ends of the axes, glabrous towards the base and moderately to densely ferrugineus pubescent towards the apex, sometimes with scattered longer silvery erect hairs; bracts broadly ovate, 35 by 35 mm, apex acute to rounded and sometimes with several narrow short teeth, ferrugineus pubescent on both surfaces, abaxial surface scattered with yellow granular glands, caducous. Pistillate flowers 34 mm long, solitary in bract axils; calyx urceolate, 12 mm long, densely ferrugineus pubescent, with scattered yellow granular glands towards the apex, persistent, splitting irregularly as ovary expands; ovary 45-carpellate, c. 1.5 mm long; styles 23 mm long, with long and finely acuminate apices, expanding as fruit matures, fused at base, free and spreading from c. 1/3 of the length, persisting to form a prominent 35 mm long crown at the fruit apex; stigma not dissected. Fruits subglobose, 68 by 1013 mm, compressed, sessile, one discrete glandular patch on each carpel wall developing into a long slender horn-like process 36 mm long, covered in yellow, sticky exudate, glabrous or rarely with some scattered ferrugineus hairs towards the base and on the sutures. Seeds subtriangular-ovoid, 44.5 mm in diam., black, with shallow coarse grooves and a small cruciform scar at the columella attachment point, completely encased in a fleshy bright red aril.
Distribution Peninsular Thailand, Malay Peninsula, Sumatra (incl. Bangka), Java, Philippines (Palawan, Sulu Prov.). Unknown for Borneo, but it has been reported from Indochina this is yet to be confirmed.
Habitat & Ecology Macaranga triloba is a small to medium sized early successional tree occurring from the lowlands up to c. 1400 m. It grows in a wide range of habitats, including dryland dipterocarp forests and on the edges of swampy forest. Macaranga triloba is one of few species in section Pachystemon that grow in seasonally dry forest; in Java it is very common on several of the mountains along the volcanic chain.
Notes 1. Thunberg (1815) did not mention a type specimen. The descriptions of Blume (1825) and subsequently Mόll.Arg. (1866) clearly refer to the same species. It seems likely that Thunberg had access to Blume's collection in writing the description, and it is therefore chosen as the lectotype.
2. There has been much misunderstanding in the application of the name Macaranga triloba. For many years the widespread ant-inhabited species, which occurs throughout western Malesia, was incorrectly referred to as M. triloba. Smith (1903) and Ridley (1910) were among the earliest biologists to remark on the extraordinary relationship formed between Macaranga triloba and ants of the genus Crematogaster. Even though Pax and Hoffmann (1914) did not describe M. triloba as having hollow ant-inhabited twigs in their revision of the genus, subsequent authors continued to refer to M. triloba as ant inhabited (Baker, 1934). An examination of the type specimens in Herbarium Bogoriense revealed that M. triloba is not in fact ant-inhabited, and that the widespread ant-inhabited species is M. bancana.
3. Macaranga triloba and M. bancana differ in numerous characteristics, and are not very closely related. In addition to having solid stems, M. triloba has erect coriaceous stipules, acuminate to caudate staminate bracteole apices, and long horns on the fruits; M. bancana has none of these features.
4. Macaranga triloba is more closely related to M. depressa f. depressa. Both species have solid stems, coriaceous stipules that are not appressed to the stem, and long horns on the mature fruits. Macaranga triloba differs in having larger leaves, with a substantially larger lamina. The leaf lobes in M. depressa f. depressa are more narrow and spreading than in M. triloba. The staminate bracteoles are larger and have a much more elongated caudate apex in M. triloba.
5. Macaranga triloba, as here circumscribed, includes the specimens previously referred to M. quadricornis Ridley. That taxon was recognized by Whitmore (1969, 1973, 1982) to occur in several disjunct populations in the Malay peninsula and in Sumatra. The observation that the taxon occurs in submontane areas in the mountains of the Peninsula and in the lowlands of Johore prompted Whitmore (1969) to infer that the species had evolved independently in these locations. However, a closer analysis of morphological variation in the species closely related to M. triloba indicates that M. quadricornis does not differ significantly from the widespread M. triloba. Macaranga quadricornis had previously been recognized on the basis of the magenta colour of the leaf undersurfaces that often (but not always) persists when the leaves mature. In other characters there are not consistent differences between these taxa.
6. Across the full ecological and altitudinal range of M. triloba there is morphological variation. For example, the acumen of the staminate bracteole appears to be particularly long in individuals from peninsular Malaysia. In addition, the degree of pubescence on the leaf undersurfaces varies substantially within that area. Further analysis of the morphological variation in this species is needed, particularly in relation to the closely related M. depressa f. depressa.
Macaranga tsonane Whitmore, Kew Bull. 34 (1980) 606; in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 160; Gen. Macaranga Prodr. (2008) 265. Type: Schodde 2454 (holo K; iso L), Papua New Guinea, Southern Highlands Dist., near Moro, Lake Kutubu.
Small trees, 10 m; entirely glabrous including innovations. Twigs 4 mm diam., very strongly angled and ridged, glabrous. Stipules ovate, 15 by 14 mm, apex acute, very strongly cucullate, dark brown papery, erect, persistent. Leaves: petioles 13 cm, weakly flattened above near base, 3 mm across, weakly kneed at base; blades ovate, 24 by 20 cm, papery, base broadly rounded 5 cm peltate, margin repand, apex acute, drying pale brown, below weakly to strongly glaucous and with close very small golden granular glands, spider's web nervation with scalariform tertiaries and quaternaries. Staminate inflorescences narrow sparsely shortly branched 22 cm racemes; peduncle 18 cm, rather flattened, 3 mm across; a few crowded divaricate 1015 mm slender branches arising from a flower cluster; bracts ovate, 8 by 4 mm, apex acute, dark brown, papery, erect, subpersistent; bracteoles ovate, 3 by 3 mm, apex 1 mm acuminate, or with several coarse 0.8 mm teeth on upper part, strongly cucullate, slightly fleshy, faintly nerved, outside glabrous, inside densely finely granular glandular, erect to spreading, enclosing flower clusters, persistent, overlapping. Staminate flowers in clusters of 34, 0.8 mm diam., sessile; stamens 3, anthers 4-locular. Fruits solitary on an 8 cm peduncle, latter flattened, 2 mm across, glabrous; fruits depressed, compressed, globose, 18 by 25 by 8 mm, blackish, densely covered with flattish tapering sinuous sparsely yellowish granular glandular 3 mm spines; pedicel stout, glabrous, 1 mm; calyx small, persistent; styles 3, free, apical, recurved, 2 mm, densely plumose. Seeds unknown.
Distribution New Guinea.
Habitat & Ecology Rain forest, sometimes sago swamp, 50780 m.
Note Highly distinctive in its cucullate stipules and long peduncled staminate inflorescence with the flowering part as a dense terminal pyramid.
Macaranga umbrosa S.J.Davies, Harvard Pap. Bot. 6 (2001) 422. Macaranga kingii Hook.f. var. platyphylla Airy Shaw, Kew Bull. 25 (1971) 533. Type: Banyeng S 27058 (holotype: K; isotypes: A, SAR, SING), Malaysia, Sarawak, Ulu Sungai Engkabang, Semengoh Forest Reserve, 12th mile Penrissen Road.
Small slender glabrous tree to 15 m tall, flowering from c. 3 m; twigs 815(18) mm in diam.r, glabrous, not glaucous, hollow, housing ants. Stipules narrowly ovate, 1527 by 813 mm, with a finely acuminate apex, recurved with the margins revolute appearing horn-like and appressed to the stem, glabrous, the pair not completely encircling the twig, succulent, producing food-bodies on abaxial surface, up to 68 pairs present on the shoots. Leaves: petioles terete, 1550 cm long, stout, up to 9 mm in diam., not glaucous, glabrous; blades ovate to broadly ovate, often huge, 2060 by 1435 cm, dark shiny green on both surfaces and appearing slightly bullate above, entire or usually with short lateral cusps, very deeply peltate to 411 cm, base broadly rounded, margin entire, with small protruding nectaries, apices acute, adaxial surface glabrous, abaxial surface glabrous or sometimes with extremely minute silvery crisped hairs on the main veins, not glaucous; 1° venation palmate with 78 prominent veins, 2° venation looping near the leaf margins and forming a subprominent marginal collecting vein and sometimes ending at the margin in small flattened nectaries, 3° venation scalariform, 4° venation loosely scalariform, finer veins loosely reticulate; young leaves dark red below, glabrous or scattered with minute ferrugineus hairs. Staminate inflorescences paniculate, erect, 615 by c. 6 cm, glabrous, up to 3 axis orders, main axis flattened, first pair of secondary branches ± opposite with or without accessory branches; bracts ovate, 815 by 510 mm, persistent, margin entire, apex acute, glabrous; flower clusters with 15 flowers, spirally arranged and crowded towards the ends of inflorescence branches; bracteoles ovate, 46 by 34 mm, thin and delicate, enclosing flower clusters, margin entire, apex acuminate, glabrous, scattered with yellow granular glands. Staminate flowers c. 1.5 mm long, pedicellate; sepals free, with scattered minute red-brown hairs and yellow glands, membranous; stamens 47; anthers 4-locular. Pistillate inflorescences particulate, erect, 312 cm, stout, glabrous, not glaucous, to 2 axis orders, secondary branches ±opposite, bracts narrowly ovate, 810 by 35 mm, glabrous, with scattered yellow glands, caducous. Pistillate flowers 815 mm, solitary in bract axils; calyx urceolate, 46 mm long, not glaucous, glabrous or with scattered ferrugineus hairs near the base, persistent, splitting irregularly as the ovary expands; ovary 45- (rarely 6)-carpellate, 46 mm long; styles 49 mm long; styles fused at base, free and spreading from c. 1/2 of the length, persisting to form a prominent 68 mm long crown at fruit apex; stigma shallowly dissected. Fruits subglobose, 912 by 1015 mm, pale pink to bright red, pedicellate, two discrete glandular patches on each carpel wall developing into narrow ridges raised to 12 mm, covered in dark, sticky exudate; pedicel 1030 mm long, glabrous, not glaucous. Seeds subtriangular-ovoid, 5.56.5 mm in diam., black, with shallow narrow grooves and a small cruciform scar, encased in a fleshy bright orange-red aril.
Distribution Borneo (Brunei and Sarawak). It is currently only known from Sarawak and Brunei, although a possible recent sighting has been reported from eastern Sabah (B. Fiala, in litt.).
Habitat & Ecology This species is a small shade tolerant tree with huge leaves on a slender stem. Macaranga umbrosa primarily occurs on relatively fertile shale-derived soils in shady or partially open areas. It is abundant on the shale-derived soils of Temburong in Brunei and the shale-derived soils of Lambir Hills National Park in Sarawak. The relatively large red fruits of this species are unusual for Malesian Macaranga; they may be more conspicuous to birds in understorey low light environments.
Notes 1. This species shares large horn-like recurved stipules, large pedicellate fruits with raised narrow ridges, and several other features with M. kingii and M. lamellata (see further discussion under M. lamellata).
2. Macaranga umbrosa differs from M. kingii in having unlobed or only slightly cusped leaves, and from M. lamellata in having glabrous leaves and not glaucous twigs.
3. Macaranga umbrosa was previously recognized as a subspecies of M. kingii. Based on the substantial ecological and morphological differences between these taxa Davies (2001) recognizes M. lamellata as a distinct species. The epithet umbrosa refers to the relatively shade tolerant habit of this species.
Macaranga uxoriae Whitmore, Gen. Macaranga Prodr. (2008) 266. Type: Versteegh BW 10276 (holo L), Indonesia, Papua, Faf Fak, Mt. Genofa.
Shrub, 8 m. Twigs 3 mm diam., terete, slightly coarsely ridged, tomentose with short white crinkled simple and tufted hairs. Stipules elliptic, 713 by 36 mm, apex acute, dark brown, papery, outside hirsute, glabrescent, erect, persistent. Leaves: pPetioles 3.56.5 cm, 2 mm diam., terete, indumentum as twigs, kneed at both ends; blades deltoid, 6.514 by 6.58 cm, thinly leathery, widest at broadly truncate very shallowly depressed base, 1 mm peltate or cordulate at petiole, with several glands on nerves above near petiole insertion, margin repand and with small 8 mm distant prominent glandular teeth, apex 1 cm acuminate to caudate, dark chocolate brown, partially weakly bullate between tertiaries, above sparsely hairy, below closely black granular glandular and velvety all nerves with short spreading simple or tufted hairs, 3 main and 2 minor nerves at petiole insertion, tertiaries and quaternaries scalariform. Staminate unknown. Infructescences dense panicles, 5.59 by 47 cm with 3 axis orders, main axis indumentum as twigs, terete, 1.5 mm diam.; peduncle 1525 mm; branches all divaricate, lowest secondaries opposite, to 3 cm long; bracts elliptic, 1.5 by 1 mm, acuminate, persistent; bracteoles round, 1.5 mm, to oblong, 2 by 1 mm, plus 1 mm caudex, with few to many adaxial patellar glands and abaxially with spreading long hairs on centre line, persistent. Fruits crowded on branches, 1 per bracteole, round, 3.5 mm across to ellipsoid, 3.5 by 3 mm, with a shallow vertical suture, leathery, black granular glandular and with sparse 1 mm glabrous tapering spines; pedicel slender, 3 mm, with sparse long spreading hairs; sepals persistent, free; style not seen. Seeds round, 2 mm, verrucose, wholly enclosed in a sarcotesta.
Distribution West New Guinea (known from type only).
Habitat & Ecology Steep slopes, rocky soil, 1000 m. Sparse in young secondary forest.
Notes 1. In combination, the characters put this into the Brunneo-floccosa group, amongst the species with bullate leaves, and collectively render it very distinctive.
2. I, slightly unconventionally, name this species in honour of my wife, Wendy. Her contribution has certainly differed from that depicted in the tailpiece (p. 566) to Stearn's Botanical Latin. In the 1960s, she typed the labels. When serious work on this Prodromus began in 1997, she became initially mistress of the database and in charge of all computing, but later has also taken on the role of assistant in all matters connected with visits to herbaria, and towards the end, also of chauffeur.
Macaranga velutina (Rchb.f. & Zoll.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 993; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 311; Whitmore, Kew Bull. 36 (1981) 322; S.J.Davies, Harvard Pap. Bot. 6 (2001) 423. Mappa velutinum Rchb.f. & Zoll., Linnaea 29 (1857) 468. Lectotype (designated by S.J.Davies, 2001): Teijsmann HB 3719 (holo: BO; iso: BO, 2 sheets), Indonesia, Sumatra, Palembang province, Oganoloe.
Macaranga depressa (Mόll.Arg.) Mόll.Arg. var. mollis Mόll.Arg. in DC., Prodr. 15, 2 (1866) 989. Type: Zollinger 4204 HB (holotype: L; isotype: BO), Indonesia, Sumatra, Lampong.
Macaranga calcicola Airy Shaw var. calcifuga Whitmore, Kew Bull. 29 (1974) 445. Macaranga calcifuga (Whitmore) R.I.Milne, Kew Bull. 49 (1994) 453. Type: Othman S 29457 (holotype: K; isotypes: KEP, L, SAN), Malaysia, Sarawak, Kuching, Semengoh Arboretum.
Macaranga indistincta auct. non Whitmore: Whitmore, Kew Bull. 29 (1974) 447, p.p.
Small tree to 15 m tall, 15 cm dbh; twigs 49 mm in diam., glabrous to densely pubescent with soft erect silvery hairs, hollow, housing ants. Stipules 68 by 69 mm, red-brown when fresh, usually finely furfuraceous when young and becoming glabrous, the pair completely encircling the twig, succulent, recurved, producing food-bodies on the abaxial surface, usually 24 pairs present on the shoots. Leaves: petioles terete, slender, (6)820 cm long, sparsely to densely covered in erect long silvery hairs; blades narrowly to broadly ovate, 1426 by 1020 cm, silvery grey to almost glaucous, tricusped to shortly trilobed or sometimes subentire, dissected to 1/81/4 (rarely more) of the leaf length, central lobe broadly acute, lateral lobes often short narrow cusps, shallowly peltate, petiole insertion 12.5 cm from the leaf margin, base broad to very broadly rounded sometimes almost truncate, margin entire, apices acute to shortly acuminate, abaxial surface densely covered in silvery-grey soft erect hairs, appearing almost glaucous; 1° venation palmate with 79 prominent veins, 2° venation scalariform and rather closely spaced, curving upwards slightly near the leaf margins and running into the margin and terminating in tiny (c. 0.3 mm) narrow conical nectaries, 3° veins densely scalariform; young leaves reddish-brown, densely covered in minute ferrugineus hairs and scattered with longer silvery hairs. Staminate inflorescences paniculate, erect, 1020 by 515 cm, densely ferrugineus tomentose towards the apex, basal axes sparsely to densely pubescent with short silvery hairs, the axes usually appearing greyish-blue, up to 4 axis orders, main basal axis flattened and less than 1/3 of the total inflorescence length, first pair of secondary branches opposite with accessory branches; bracts ovate, 35 by 23 mm, margin entire, apex acute, finely ferrugineus pubescent on both surfaces and the margin, caducous; flower clusters with 1525 (30) flowers, spirally arranged and scattered along ultimate inflorescence branches; bracteoles small, broadly ovate to transversely elliptical, 1.52.5 by 24 mm, enclosing flower clusters, margin entire, apex rounded or often with a shortly acute tip, densely covered with minute ferrugineus hairs and yellow granular glands. Staminate flowers c. 0.7 mm long, sessile; sepals fused, splitting irregularly to c. 1/3, apex densely covered in minute red-brown hairs; stamens 1; anthers 3-locular. Pistillate inflorescences particulate, erect, 38(11) by 35(7) cm, stout, sparsely to densely covered in short greyish hairs, up to 3 axis orders, secondary branches opposite and usually with accessory branches; bracts elliptic, 58 mm long, c. 3 mm wide at the base, with an acute apex, adaxial surface with scattered hairs, caducous, abaxial surface scattered with minute silvery-grey hairs and tiny yellow granular glands particularly towards the base. Pistillate flowers c. 2 mm long, solitary in bract axils; calyx urceolate, 11.5 mm long, with scattered ferrugineus and minute silvery hairs, persistent, splitting irregularly as ovary expands; ovary 4-(rarely 5-)carpellate, c. 1.5 mm long; styles c. 1 mm long, fused at base, free and spreading from c. 1/2 of the length, persisting to form a prominent 23 mm long crown at fruit apex; stigma not dissected. Fruits subglobose, compressed, 57 by 1011 mm, greyish-blue green and slightly reddish on the apex when fresh, sessile, glabrous, with one discrete glandular patch on each carpel wall developing into a short pointed swelling c. 12 mm long, covered in yellow-green, sticky exudate. Seeds subtriangular-ovoid, c. 4 mm in diam., black, with shallow coarse grooves and a small cruciform scar at the columella attachment point, encased in a fleshy bright pinkish to orange-red aril.
Distribution Sumatra (incl. Bangka), Borneo (Kalimantan and Sarawak). Macaranga velutina is not known from northwestern Sarawak, Brunei or Sabah. In these areas it appears to be replaced by the ecologically very similar species, M. bancana and M. indistincta.
Habitat & Ecology An early successional tree common in degraded forest, particularly in wet areas, such as in road side gullies, along the margins of peat swamp forest and in abandoned rubber plantations.
Notes 1. The original description of Mappa velutinum Reichenbach & Zollinger (1857) cite a Teijsmann collection from Oganulu [sic] from the Province of Palembang. The only Teijsmann collection S.J.Davies (2001) has seen from that location is numbered 3719. There are three sheets of this in BO, one of which indicates the specific location and is designated the lectotype.
2. The original description Macaranga depressa var. mollis Mόll.Arg. does not cite a Zollinger number but the collection Zollinger 4204 HB is from Lampong. It is also cited by Whitmore in Kew Bull. 29 (1974) 447.
3. Macaranga velutina is an ant-inhabited species of section Pachystemon that is closely related to M. bancana and M. indistincta. All three species are early successional small to medium-sized trees that are common along roadsides and in degraded land. The distributions of these three species do not overlap, with the exception of M. bancana and M. velutina which are both known from the island of Bangka. Field work in Bangka is required to assess whether these species are locally sympatric.
4. Macaranga velutina and M. indistincta both have densely pubescent lower leaf surfaces, whereas the lower leaf surface of M. bancana is glabrous or has scattered minute hairs on the veins only. Macaranga velutina differs from M. indistincta in having entire staminate bracteoles, and fruits with very short horns. Macaranga velutina is probably more closely related to M. bancana; both species have small entire staminate bracteoles, and fruits with short horns. In addition to having pubescent leaves, Macaranga velutina differs from M. bancana in having generally less deeply-dissected leaves, bluish to almost glaucous inflorescences with silvery hairs towards the base, and young stems that are greyish-blue to almost glaucous.
5. There is some morphological variation in M. velutina across its geographical range. In Sarawak and east Borneo it has glabrous and somewhat glaucous twigs, whereas the material from central and west Borneo, Sumatra and Bangka is densely pubescent on the twigs. Further work on this species complex is required.
Macaranga velutiniflora S.J.Davies, Harvard Pap. Bot. 4 (1999) 433, Fig. 1; Harvard Pap. Bot. 6 (2001) 425. Type: Davies 981201 (holotype: SAR; isotype: A), Malaysia, Sarawak, Padawan area, just below entrance to Puncak Borneo tourist resort, c. 800 m.
Tree 815 m tall, c. 15 cm in diam., with huge pendulous leaves held in a shallow canopy at the stem apex; twigs very stout, terete, 814 mm in diam., glabrous, not glaucous, housing ants. Bark smooth, light brown, hoop-marked. Stipules massive, deltate, 1114 by 914 mm, apex acute, pale green to reddish-brown when fresh, recurved, succulent, glabrous, completely encircling the twig, with 24 pairs present on the shoots, producing food-bodies on the abaxial surface. Leaves: petioles terete or sometimes slightly flattened near base, stout, 1228 cm long, glabrous; blades very large, very broadly ovate, 2560 by 2545 cm, peltate, petiole insertion >6 cm from the margin, shallowly trilobed to tricusped, dissected to 1/101/5 of the leaf length, base broadly rounded to almost truncate, margin entire with secondary and tertiary veins terminating in c. 1 mm diameter protruding flattened nectaries, apices broadly acute to shortly acuminate, lamina mostly glabrous but adaxial venation with scattered long hyaline hairs and abaxial venation with very minute scattered tomentum, not glaucous, 1° venation palmate with 1011 prominent veins, minor veins prominent and densely scalariform, 2°4° veins densely scalariform; young leaves reddish-brown, with short silvery velutinous hairs and scattered erect spiny hairs, soon becoming glabrous. Staminate inflorescences paniculate, erect, spreading, very broad, 1535 by 1530 cm, covered in a dense velvety indumentum of short thick pale golden-brown hairs on all axes except the basal rhachis, main axes also sparsely to densely covered in long erect hairs, not glaucous, densely branched with up to 5 axis orders; main basal axis flattened, 2.59 cm long; secondary branches opposite with accessory branches; bracts narrowly triangular, 512 mm long, caducous, margin entire and fringed with minute silvery hairs, apex acute, adaxial surface mostly glabrous, abaxial surface finely pubescent near base and along midrib; flower clusters of 1530 flowers alternate and scattered along inflorescence branches; bracteoles broadly ovate to sub-orbicular, 23 by 24 mm, enclosing the flowers, margin entire, apex rounded to acute, abaxial surface with 610 subprominent ribs and yellow granular glands, both surfaces densely velvety pubescent with pale golden-brown stout hairs. Staminate flowers c. 1 mm long, with dense silvery hairs on the apex, sessile; sepals fused, splitting; stamens 1; anthers 3-locular. Pistillate inflorescences particulate, erect, 514 by 613 cm, stout, up to 3 axis orders, covered in a dense velutinous indumentum of thick pale golden-brown hairs on all axes except the main basal rhachis, main axes densely covered in long erect hairs; basal unbranched axis flattened and 14 cm long, first pair of secondary branches opposite with accessory branches; bracts narrowly triangular, c. 9 mm long, adaxial surface velutinous towards the base and along the midrib, otherwise glabrous, adaxial surface ± glabrous with some long villous hairs towards the base, caducous. Pistillate flowers 45 mm long, solitary in bracteole axils, sessile; calyx urceolate, 23 mm long, densely silvery to golden velutinous with slightly longer hairs towards the apex, persistent, splitting irregularly; ovary c. 3 mm long, (4)5(6)-carpellate, densely velutinous along sutures and towards base, each carpel with one discrete patch of dense yellow granular glands on the surface; styles 1.52 mm long, fused at the base, free and spreading from 4/5 of length, persisting to form a broad c. 2 mm long crown at the apex of the fruit; stigma not dissected. Fruits (not quite mature) subglobose, 811 by 1214(16) mm, sessile, finely velutinous on the sutures and base, glandular patch on each carpel wall developing into a long horn-like process covered in a sticky yellow-green glandular exudate. Seeds subtriangular-ovoid, 44.5 mm in diam., colour not seen, surface finely grooved and pitted, with small cruciform scar, completely encased in a fleshy aril (colour not seen).
Distribution Borneo (West Kalimantan and Sarawak). It has been collected from two locations: the Padawan area near Gunung Penrissen in Sarawak, and Gunung Palung in West Kalimantan, Indonesia.
Habitat & Ecology Macaranga velutiniflora is an early successional tree of primary and disturbed mixed dipterocarp forest. The Sarawak collections are from c. 800900 m above sea level and the Kalimantan collections are from the lowlands. Further exploration is required to assess the ecological range of this species.
Notes 1. Macaranga velutiniflora is a member of the largest clade of section Pachystemon. This species appears to be most closely related to another submontane species, M. petanostyla Airy Shaw, or to the lowland species, M. trachyphylla Airy Shaw. These three species share several morphological features including: large and shallowly trilobed leaves, very stout twigs with exceptionally large succulent recurved stipules, and long horn-like processes on the fruit. Macaranga velutiniflora differs from these species in the velvety pubescence of the staminate and pistillate inflorescences, and in having broadly ovate staminate bracteoles with rounded to shortly acute apices and several subprominent parallel ribs on the abaxial surface.
2. The two staminate collections (Laman 1181 and Laman 630) from Gunung Palung in West Kalimantan differ slightly from the Sarawak collections. While the Kalimantan specimens are very similar in most details of the leaves, twigs and inflorescences, they differ from the Sarawak collections in having darker red-brown velutinous pubescence on the inflorescence axes as compared to pale golden brown hairs in the Sarawak specimens. In addition, the bracteoles of the Kalimantan collections do not appear to be at all ribbed on the abaxial surface as in the Sarawak collections. Finally, the Kalimantan collections are from 40 m elevation whereas the Sarawak collections are from 800900 m. Since the staminate Kalimantan collections match the staminate Sarawak collections in most respects, they are included under M. velutiniflora. Further work is required to clarify whether they may in fact represent a different species.
Macaranga versteeghii L.M.Perry, J. Arnold Arb. 34 (1953) 242; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 160; Gen. Macaranga Prodr. (2008) 269. Type: Brass & Versteegh 11116 (holo A; iso L), Indonesia, Papua, Lake Habbema, Bele River.
Trees, 515 m. Twigs 3 mm diam., slightly flattened, densely rusty brown pubescent at tip. Stipules oblong-elliptic, 5 mm, apex acute or sub-acuminate, velvety, rigid, soon caducous. Leaves: petioles 4.5 cm, 1.5 mm diam., terete, ridged, at first densely rusty brown pubescent; blades broadly ovate, 7.516 by 49.5 cm, leathery, base rounded, 5 mm peltate, most with 1few small inconspicuous glands near base, margin slightly concave at base, weakly repand, apex abruptly narrowing to 1 cm tip, drying mid brown, glabrous above, below weakly pubescent on main nerves and sparsely granular glandular, secondaries and reticulations slightly raised above, strongly raised below, slightly irregularly bullate between some mainly lower secondaries, with 3 main and several minor nerves from petiole insertion, secondary nerves including basal only 57 pairs. Staminate inflorescences (young) 57 cm rusty tomentose panicles; bracteoles mostly elliptic, 3.5 mm, a few broadly ovate, 1.5 mm, abruptly and briefly acuminate. Staminate flowers buds tomentellose and granular glandular; stamens 914, stamens 4-locular. Infructescences 8 cm spikes, or racemes with sparse sinuous spreading 4 cm branches; peduncle 3 cm; main axis slightly flattened, rusty pubescent; bracts ovate, 1 mm, apex blunt, thick, pubescent, persistent; bracteoles not seen. Fruits solitary or paired, irregularly spaced along secondary axes, roundish but slightly angled, weakly bilobed, 5 mm diam., leathery, with sparse small spiny tubercles (spines brittle), evanescent scurf and sparse granular glands; pedicel absent or stout, 2 mm; calyx persistent; style single, 5 mm, sub-basal, minutely papillose, caducous. Seeds (young) roundish, tuberculate, with sarcotesta.
Distribution West New Guinea (restricted to Baliem valley, Mt.Habbema area).
Habitat & Ecology Primary or secondary montane forests, 20002500 m.
Note Stipules and staminate described mainly from Perry (1953). Her panicles' may be racemes.
Macaranga villosula Pax & K.Hoffm. in Engl.,Pflanzenr. IV.147.xiv add. vi (1919) 27; L.M.Perry, J. Arnold Arb. 34 (1953) 249; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 160; Gen. Macaranga Prodr. (2008) 270. Syntypes: Ledermann 9787 (K, L), Papua New Guinea, Sepik Region, April River; Ledermann 10593 (not seen), idem, Malu.
Trees, 412 m tall. Twigs 3 mm diam., angled, pubescent or furfurescent. Stipules ovate, to 5 by 3 mm, apex acuminate, dark brown, papery, variously tomentose, erect, caducous. Leaves: petioles 5 cm, 2 mm diam., terete, sparsely to densely pubescent, kneed at top; blades ovate, c.18 by 8cm, papery, base rounded with 23 small protruding glands at petiole insertion, margin repand, apex abruptly 15 mm acuminate, drying mid brown, sparsely to densely hairy on midrib and main nerves below, granular glands dark sparse to fairly dense. Staminate inflorescences 6 cm pubescent racemes; peduncle very short; main axis slender; branches thread-like (?); bracteoles small. Staminate flowers in clusters of 56, shortly pedicellate; calyx pubescent; stamens 3, anthers 4-locular. Pistillate inflorescences mostly behind leaves, 6 cm densely pubescent racemes; peduncle 1 cm; main axis flattened, 1.5 mm; branches spreading, alternate or opposite, thread-like; bracteoles ovate, 1 by 0.5 mm, apex acute, outside pubescent, spreading, persistent. Pistillate flowers: sepals 3, triangular, 0.5 mm long, apex acute; ovary 1-locular, unarmed, densely granular glandular, style 4 mm, slender, papillose. Young Fruits solitary, thinly woody, black granular glandular, sessile; calyx persistent; style subbasal, 1 cm, thread-like, sparsely papillose, subpersistent; at maturity 3 mm diam., sessile or pedicel very short; style lost. Seeds globose, finely verrucose, with sarcotesta.
Distribution Moluccas (Aru, Kobrool), East New Guinea.
Habitat & Ecology Primary forest to 400 m. Rare.
Note The stamiante part is described here from the original description. All other species of the Angustifolia group have the stamiante bracteoles smaller than the flower clusters, so this probably has. If so, that is a good distinction from the otherwise similar M. polyadenia. The pistillate flowers are also described from the original.
Macaranga warburgiana Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 347; L.M.Perry, J. Arnold Arb. 34 (1953) 215; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 160; Gen. Macaranga Prodr. (2008) 271. Syntypes: Hellwig 403, Nyman 680, Warburg s.n. (all not seen), Ppua New Guinea, Morobe Prov., Sattelberg, Finschhafen.
Macaranga cuspidata Warb., Bot. Jahrb. 13 (1891) 351; K.Schum. & Lauterb., Fl. Schutzgeb. Sόdsee (1900) 396 (non Boiv. ex Baill., Adansonia 1, 18612: 260).
Small trees, to 5 m. Twigs strongly angled, 3 mm diam., usually blackish brown with pale stipule scars, glabrous. Stipules oblong-elliptic 1215 by 4 mm, apex acuminate, blackish brown, papery, glabrous or with a few hairs or weakly granular glandular, erect, usually soon caducous. Leaves: petioles c.8 cm, slender, 2 mm diam., terete, smooth, at first with sparse white spreading hairs, soon glabrous, weakly kneed at both ends; blades ovate, 923 by 913 cm, papery, base broadly shallowly cordate to broadly rounded or broadly truncate, with several roundish glands scattered near petiole insertion above, margin often coarsely distantly glandular dentate, apex cuspidate, 2 cm, drying khaki with on upper side main nerves usually darker, glabrous, 3 main nerves at petiole insertion. Staminate inflorescences slender panicles with 3 axis orders, to 9 cm long, glabrous, branches divaricate or ascending; peduncle to 7 cm, flattened, 2 mm across, striate, with 57 thread-like 3 cm secondary branches all arising from a flower cluster, subtended by a sometimes persistent leafy elliptic bract to 17 by 3 mm including 2 mm stalk and 5 mm caudex, papery, pinnately nerved, granular glandular; secondary branches with a few 5 mm tertiary branches and 3 mm spaced small flower clusters subtended by spreading or upturned at tip bracteoles longer than cluster, oblong, 2 by 1 mm, apex acute, slightly narrowing towards base, margin entire, with 2 adaxial patellar glands. Staminate flowers 4 per cluster 0.6 mm diam., sessile; sepals glabrous with a few granular glands; stamens 56, anthers 4-locular. Pistillate inflorescences very long spikes to 26 cm; axis terete, 2 mm diam., sparsely hirsute with long white hairs, becoming glabrous, with terminal and subterminal clusters of 24 fruits subtended by caducous elliptic leafy bracts, 16 by 3 mm cucullate at base, with 8 by 1 mm narrow acuminate tip, glabrous, with scattered small adaxial patellar glands, margins entire with a few small protruding marginal glands. Fruits mostly shallowly bilobed, 5 by 9 mm, sparsely shortly spiny on upper parts of lobes away from suture, granular glandular, woody, sessile with persistent calyx. Seeds round, 5 mm, with sarcotesta, rugose.
Distribution East New Guinea, Bismarck Archipelago (New Ireland).
Habitat & Ecology Secondary forest and primary forest edges, hills and mountains, 2701350 m.
Note For the identity of this species see discussions in Perry (1953) and Whitmore (1980). The very unusual stamiante inflorescence is described mainly from Frodin UPNG 4172.
Macaranga waturandangii Whitmore, Gen. Macaranga Prodr. (2008) 272. Type: Waturandung 49 (holo K; iso BO), Sulawesi, Malili.
Twigs 5 mm diam., terete, blackish, almost glabrous, weakly rusty furfuraceous. Stipules elliptic, 5 by 1 mm, rusty furfuraceous, erect, soon caducous. Leaves: petioles slender, 8 cm, terete, sparsely rusty furfuraceous, not kneed; blades ovate, 1116 by 7.513 cm, thinly leathery, base broadly cuneate to truncate, sometimes minutely 12 mm peltate, without glands near insertion, margin entire, apex long acute, drying grey green above, rusty brown below, nerves and reticulations below finely pubescent, tertiary nerves conspicuous scalariform. Staminate inflorescences behind the leaves, dense panicles to 20 by 6 cm, 3 axis orders, finely rusty furfuraceous; branches spreading; peduncle to 5 cm; main axis stout, terete, somewhat ridged; branches alternate, without accessories; bracts soon caducous, not seen; ultimate branches thread-like, bearing flower clusters, crowded at first, ultimately evenly spaced; bracteoles persistent, ovate, 2 by 1.5 mm, apex acute, upper margins coarsely dentate, without patellar glands, both sides furfuraceous, erect, greater than flower cluster. Staminate flowers 6 or more per cluster, very tiny, 0.3 mm diam., sessile; sepals finely furfuraceous outside; stamens 34, anthers 4-locular. Pistillate inflorescences as staminate but narrow racemes to 10 cm. Fruits in distant groups of 23, bilobed, depressed globose, 3 by 4 mm, thinly woody, with sparse granular glands; pedicel 5 mm, slender, furfuraceous; calyx peristent; styles 2, recurved, caducous, 1 mm, apical. Seeds coarsely shallowly rugose, with sarcotesta.
Distribution Sulawesi.
Macaranga winkleri Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 355; Mitt. Inst. Bot. Hamburg 7 (1931) 228; Whitmore, Gen. Macaranga Prodr. (2008) 275, Fig. 25. Type: Winkler 2873 (not seen), Borneo, Kalimantan, Batu Babi to Lumowia.
Slender tree to 22 m tall, though often flowering from c. 5 m. Bark smooth, remaining green at maturity, often with prominent ant holes. Twigs glabrous, hollow, 714 mm diam., housing ants, with prominent ant holes often in narrow grooves. Stipules narrowly ovate, 1525 by 813 mm, with a long finely acuminate apex, glabrous, coriaceous, not succulent, recurved with the lateral sides appressed and appearing horn-like, producing food-bodies on the abaxial surface, numerous pairs present on the shoots, the pair not completely encircling the twig. Leaves: petioles terete, 1635 cm long, slender, glabrous; blades ovate, 1640 by 1225 cm, base rounded, 38 cm peltate, margin entire, with small protruding conical glands, leaf apex acute to finely acuminate; adaxial and abaxial surfaces entirely glabrous, not glaucous, patellar glands absent from lamina; 1o venation palmate, with (5)67(8) prominent veins, 2o venation looping just inside the leaf margins, 3o venation scalariform, 4o venation weakly scalariform; leaf margin entire; young leaves glabrous, characeous, abaxial surface densely punctate glandular. Staminate inflorescences paniculate, erect, 2035 by 1220 cm, glabrous, 4 axis orders, main axis terete or slightly angular when dry, first pair of secondary branches alternate without accessory branches; bracts narrowly ovate, 1025 by c. 6 mm, apex acute, persistent, base often slightly cordate, margin entire, glabrous, usually reddish; flower clusters with 615 flowers, spirally arranged, evenly spaced forming catkin-like clusters on the ultimate inflorescence branches; bracteoles ovate, 25 by 1.54 mm, saccate, enclosing flower clusters, apex rounded, margin deeply dissected into 48 slender 23 mm long tentacular lobes with small conical apical glands at apices, abaxial surface with several subprominent ribs and scattered to densely covered with yellow granular glands, otherwise glabrous, adaxial surface densely ferruginous pubescent particularly towards the base. Staminate flowers c. 1 mm long, shortly pedicellate; sepals free, apex with scattered yellow glands, glabrous or with a few ferruginous hairs at base; stamens 23; anthers 4-locular. Pistillate inflorescences paniculate, erect, 1026 by 815 cm, glabrous, to 3 axis orders, secondary branches alternate, glabrous or with scattered minute ferruginous hairs towards apices; bracts ovate, c. 1 cm long, often slightly cordate, usually persistent; bracteoles ovate, c. 3 by 2 mm, with 36 short narrow gland-tipped lobes, abaxial surface densely yellow glandular, adaxial surface densely ferruginous pubescent. Pistillate flowers 13 mm long, 25 in bract axils, pedicellate; calyx urceolate, c. 1 mm long, with minute yellow granular glands, glabrous, persistent and spreading at base of fruit; ovary 2-carpellate, c. 1 mm long; styles short and thick, c. 0.8 mm long, broad, free, persistent; stigma not dissected. Fruits 22.5 by 34 mm, bilocular, locules globose, pedicellate, carpel walls with 48 minute knob-like caducous processes, evenly covered in yellow granular glands, sticky; pedicel 37 mm long, slender, with yellow granular glands, glabrous. Seeds c. 1.5 mm diam., ovoid, black, very coarsely verrucose, with a small cruciform scar, encased in a fleshy violet aril.
Distribution Borneo.
Habitat & Ecology Slik et al.: In disturbed sites (gaps) in primary forest and in secondary forests, including very disturbed forest (after fires). Often along forest edges, road sides, river sides, in land slips. On yellow sandy loam. Usually ant inhabited. Up to 1800 m altitude.
Macaranga winkleriella Whitmore, Kew Bull. 29 (1974) 449; Gen. Macaranga Prodr. (2008) 277. Type: Chew Wee Kek 45y (holo K), Borneo, Sarawak, Baram Distr., Melinau Gorge.
Small trees or treelets, 1.54.0 m tall. Twigs 57 mm diam., glabrous, sometimes slightly glaucous, hollow, housing ants. Stipules broadly ovate, 46 by 46 mm, with a rounded or shortly acute apex, recurved, coriaceous, not succulent, producing food-bodies on abaxial surface, usually many pairs (28) present on the shoots. Leaves: petioles terete, reddish, 1020 cm long, sometimes slightly glaucous, glabrous; blades ovate, 1220 by 812 cm, base rounded, 2.54.0 cm peltate, leaf margin entire, apix finely acuminate, both surfaces entirely glabrous, not glaucous; young leaves red-brown, glabrous, without patellar glands; 1o venation palmate with 78 prominent veins, 2o venation scalariform with veins looping near the leaf margin and sometimes ending in small protruding conical glands, 3o venation scalariform, finer nerves loosely reticulate. Staminate inflorescences shortly paniculate, erect, 2.55 by 12.5 cm, glabrous, 3 axis orders, with flowers often produced around the base of inflorescence branches; main axis flattened; first pair of secondary branches + opposite without accessory branches; bracts deltate, c. 7 by 4 mm, caducous, apex acute, margin entire, glabrous; flower clusters with 1015 flowers, spirally arranged and evenly spaced on inflorescence branches; bracteoles ovate to obovate and slightly narrowed towards the base, c. 3 by 2 mm, erect, apex rounded with margin dissected to about 1/2 of the length into (4)5 spreading narrow flat lobes (sometimes gland-tipped), each lobe having a slightly raised subprominent vein on the abaxial surface, completely glabrous on both surfaces, rarely with a few scattered tiny ferruginous hairs towards the base. Staminate flowers c. 0.75 mm long, pedicellate with the pedicels often persisting after the flowers fall; sepals free, apex glabrous with scattered yellow granular glands; stamens 34; anthers 4-locular. Pistillate inflorescences shortly paniculate, erect, 26 cm long, 1.54 cm wide, slender, glabrous, sometimes glaucous, 2 orders of branching, often with flowers produced around the base of inflorescence branches, first pair of secondary branches alternate; bracteoles soon caducous, deltate, 22.5 mm long, apex acute to acuminate, margin entire or sometimes dissected with 23 narrow spreading elongage lobes but not as dissected as in the staminate bracteole. Pistillate flowers c. 2 mm long, in clusters of 13 in bracteole axils; calyx urceolate, enclosing the ovaries, c. 1 mm long, apex glabrous with scattered yellow granular glands, persistent, splitting irregularly as ovary expands; ovary 2-carpellate, c. 1 mm long; styles 1.52 mm long, free and spreading, persisting at fruit apex; stigma not dissected. Fruits c. 3 by 4.55.5 mm, bilocular, locules globose, pedicellate, carpel walls each with 46 tiny caducous knob-like processes, evenly covered in yellow granular glands producing sticky exudate; pedicel 46 mm long, slender, glabrous. Seeds 22.5 mm diam. (not fully mature), ovoid, covered in shallow grooves, with a small, rather faint cruciform scar, encased in a fleshy aril.
Distribution Borneo (Sarawak).
Note All collections appear to have been on limestone, although only two collections listed this.
Macaranga yakasii Airy Shaw, Kew Bull. 32 (1978) 410; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980) 161; Gen. Macaranga Prodr. (2008) 278. Type: Croft & Lelean LAE 65615 (holo K; iso L), Papua New Guinea, New Ireland, Kavieng Subdistr., Schleinitz Range, 5 km S of Logagon village.
Trees, 830 m tall, to 50 cm diam. Twigs stout, 15 mm diam., near tip, 23 mm diam. at base, terete, with conspicuous petiole scars, densely tawny pubescent, with some long hairs. Stipules elliptic, 2550 by 510 mm, tapering strongly from a broad base, apex acuminate, outside tawny pubescent, inside glabrous, often leathery, conduplicate, long persistent, erect, the upper ones forming a tuft protruding beyond apical bud. Leaves: petioles c. 40cm, stout, 1 cm diam. at base, weakly flattened above below, terete upwards, coarsely shallowly striate, tawny furfuraceous, at first densely so, base drying slightly contricted, apex weakly kneed; blades big, ovate, to 60 by 44 cm, rather papery, base broadly rounded, deeply 912 cm peltate, margin repand, apex regularly acuminate or broadly 2 cm caudate, drying chestnut to greyish brown above, below tawny with lamina surface lightly glaucous, above glabrous except main nerves near base tawny pubescent, below all nerves with spreading long white tufted hairs and variously fine tawny pubescent, close tiny black granular glands, spider's web nervation, tertiaries and quaternaries raised below, regularly scalariform. Staminate inflorescences rather open 19 by 8 cm sinuously branched panicles with 3 axis orders, with evanescent tawny pubescence; secondary axes ascending often adpressed to main axis, to 8 cm, flattened, slender, some with axillary branches(?); peduncle c. 15 mm (?), flattened, 3 mm diam. (?), ridged, narrowing upwards into main axis; branches alternate; bracts a few persistent, strap-like narrowly oblong, to 10 by 2 mm, apex blunt, dark brown papery, evanescent tawny pubescent; tertiary axes few, to 1cm, branches arising from within the puberulous flower clusters; latter grouped (especially at tips) or to 1 cm spaced; bracteoles flange-like, 2 by 3 mm, apex blunt or very broadly acute, spreading, persistent, (thinly) fleshy, smaller than cluster, glabrous with adaxially 12 inconspicuous fairly large patellar glands, margin entire. Staminate flowers 50 in tight 4 mm diam. clusters, tiny, obconical, 3 mm long, 1 mm at truncate tip; calyx 5-lobed, pubescent; stamens 3, anthers 4-locular. Infructescences 610 cm long, stout, puberulous racemes; peduncle angled, 2 cm, woody, 24 mm diam.; branches divaricate, 3 cm long, tawny. Fruits single or paired, large, depressed, squarish, 9 by 9 by 14 mm, with 4 conspicuous sutures, and slightly indented lobes, rarely 3-lobed, woody, surface irregularly with rather sparse short stout subacute evanescently tawny processes and below that obscurely granular glandular; pedicel very stout, 23 mm long, 1 mm diam.; calyx tiny, fused, tawny, reflexed, 2 mm diam., persistent; styles 34, stout, reflexed, persistent, 3 mm, tawny, densely shortly plumose. Seeds globose, 6 mm, distantly regularly verrucose, with sarcotesta.
Distribution Bismarck Archipelago (New Ireland).
Habitat & Ecology Lowlands, swamps, dryland, sometimes karst.
Note The staminate inflorescence structure cannot be seen in full on the only staminate sheet seen (Coode et al. LAE 29730, K!).
Macaranga brachythyrsa Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 320; Whitmore, Gen. Macaranga Prodr. (2008) 280. Type: Winkler 3319 (not seen), Indonesia, Borneo, Djihi.
Note Pax & Hoffmann (1914) placed this species in section Semiglobosae, but the type has so far not been located (S.J. Davies, pers. comm.).
Mappa cissifolia Rchb.f.& Zoll., Linnaea 29 (1857) 466. Macaranga cissifolia (Rchb.f. & Zoll.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 993; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 392; Davies, Harvard Pap. Bot. 6 (2001) 440; Whitmore, Gen. Macaranga Prodr. (2008) 280. Type: Teijsmann s.n., 1857 (not seen), Sumatra, Bangka. See note, also under M. cuneifolia.
Description Twigs hollow, pruinose. Stipules broadly emarginate, mucronate, glabrous, reflexed-adpressed. Leaves: petioles longer than blades; blades outline rounded, deeply cordate, deeply trilobed, lobes acuminate, repando-dentate, central lobe obovate, laterals oblique, lower part of inner margins entire, nerves subvillous, below punctate and finely villous.
Distribution Sumatra (Bangka)
Notes 1. I think the type of this is as listed above, see notes under M. cuneifolia.
2. The description is translated from Zollinger's latin in Linnaea. Until this sterile collection is found (which is unlikely) M. cissifolia cannot be placed. It resembles M. hosei in its pruinose stem and cordate leaves subvillous below, though that has spreading not reflexed-adpressed stipules, and was published in 1857, not 1887, so is an earlier name.
Adelia cordifolia Roxb., Fl. Ind., ed. 2, 3 (1832) 849. Macaranga cordifolia (Roxb.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1002; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 392; Airy Shaw, Kew Bull. 37 (1982) 26; Whitmore, Gen. Macaranga Prodr. (2008) 280. Type: see Note.
Description Leaves: petioles long; blades round, cordate, entire. Staminate spikes axillary, anthers with 4 loculi.
Distribution: Moluccas.
Note Roxburgh's (1832) description, as paraphrased by Mόller (1966), see above, is inadequate. Forman (Kew Bull. 52, 1997, 573574) in his analysis of Roxburgh's material found no material or plate. Mόller (1866) made the transfer to Macaranga, but neither he nor Pax & Hoffman saw any material. As Airy Shaw (1982) commented: quid? We shall probably never know what Roxburgh was describing.
Macaranga eloba Pax & K.Hoffm., Mitt. Bot. Mus. Hamburg 7 (1931) 228; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 159; Gen. Macaranga Prodr. (2008) 281. Type: Winkler 968a (not found), Indonesia, West Kalimantan, Bukit Raya. See Davies, Harvard Pap. Bot. 6 (2001) 440.
This species was described to have entire peltate leaves, stout twigs with short pilose hairs soon becoming glabrous, ovate-elliptic reflexed stipules with acuminate apices, short (c. 4 cm long) infructescences, and 4-locular sessile fruits bearing four 1 cm long horns. This species is clearly a member of the largest clade of section Pachystemon. Davies (Harvard Pap. Bot. 6, 2001, 438) suspects that it may either be M. hullettii or M. rostrata. Both species are known from above 1200 m altitude, have entire leaves, and have long horn-like appendages on the fruits. The stipule description sounds more like M. hullettii, but there is no mention in the description as to whether the twigs are hollow and ant-inhabited. It would be useful to relocate this specimen prior to a detailed reassessment of the morphologically variable M. hullettii. Whitmore (1975) suggested M. eloba may be M. caladiifolia; this is unlikely since that species has two short ridges on each carpel and not 1 cm long horns.
Macaranga gossypifolia Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 311; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 4 (1975) 159; S.J.Davies, Harvard Pap. Bot. 6 (2001) 440; Whitmore, Gen. Macaranga Prodr. (2008) 281. Type: Winkler 2996 (not found), Indonesia, Kalimantan, between Simpokak and Semurung. See Davies , Harvard Pap. Bot. 6 (2001) 440.
The species is described as having stout glaucous twigs, deeply evenly dissected trilobed leaves with a deeply cordate base and villous hairs on the abaxial surface, broadly triangular-ovate stipules (7 by 10 mm), and pectinate-dentate bracteoles. The deeply cordate leaves suggest that this specimen may be M. hosei, although it is not mentioned whether the leaves are peltate or not.
Rottlera hexandra Roxb., [Hort. Beng. (1814) 105, nom. nud.] Fl. Ind. ed. 2, 3 (1832) 829 ('hexandria'). Macaranga hexandra (Roxb.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1003; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 393; Whitmore, Gen. Macaranga Prodr. (2008) 281. Type: not indicated, Malay Islands.
Macaranga parvibracteata Pax & K.Hoffm. in Engl.,Pflanzenr. IV.147.xiv add. vi (1919) 24; Airy Shaw, Kew Bull. 37 (1982) 27; Whitmore, Gen. Macaranga Prodr. (2008) 281. Type: Warburg 17280 (not seen), Moluccas, Ambon.
Mappa teysmannii Zoll., Linnaea 29 (1857) 465 (teyssmannii'). Macaranga teysmannii (Zoll.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 999; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 339; Whitmore in Airy Shaw, Kew Bull. 36 (1981) 323; Gen. Macaranga Prodr. (2008) 281. Type: Teijsmann s.n., 1857 (not found), Sumatra, Palembang, Kebu Lahat.
Description Twigs hollow, subpruinose, pale pubescent at the tip. Stipules oblong, apex obtuse, keeled, erect, glabrous. Leaves: petioles long, pubescent at apex; blades rotund-ovate, peltate, apex acute, upper margins sinuous, glabrous, granular glandular below. Staminate inflorescences crowded racemes, scaly (sic!), glabrescent. Pistillate flowers solitary, calyx urceolate with 2 short thick recurved stigmas, ovary peltate-glandular.
Distribution Sumatra.
Note Zollinger's (1857) description translated here is inadequate to identify this species, based on an un-numbered Teysmann collection. He compares it to M. tanarius but the hollow twigs and female flower description suggest the Hypoleuca group. How he described staminate and pistillate parts from a single collection is a mystery. This species will remain incertae sedis until a sheet annotated by Zollinger is found.
Mappa cuneifolia Rchb.f. & Zoll., Linnaea 29 (1857) 467. Macaranga cuneifolia (Rchb.f. & Zoll.) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 1004; Pax & K.Hoffm. in Engl.,Pflanzenr. IV.147.vii (1914) 392; Whitmore in Airy Shaw, Kew Bull. 36 (1981) 322. Type: see note.
Twigs angular, velutinous. Leaves: petioles velutinous; blades rhomboid-ovate or ovate, base broad or frequently cuneate, margin entire and upwards doubly sinuous with subulate glandular teeth, apex acuminate, velutinous above, softly tomentose below. Staminate inflorescences shorter than the petioles, erect, bracteoles boat-shaped, apex acuminate.
Distribution Sumatra (Palembang, G. Batu: Herb. R.Z. no 1760, W!).
Notes 1. This description, translated from the latin description of Reichenbach and Zollinger (1857) in Linnaea, is a good match for the sheet at W listed above, whose label is in Zollinger's handwriting, and which also says Mappa cuneifolia RZ. I assume from this evidence that this must be the type of Mappa cuneifolia. Unfortunately, in Linnaea this number (as Herb. R.Z. no 1760') is cited as the type of Mappa cissifolia, a totally different, trilobed species. The type of Mappa cuneifolia is cited in Linnaea as Insula Banka ?, cl. Teyssmann' [note the misspelling]. I think Zollinger made a mistake in Linnaea, switching these two collections. This mistake has been copied by all subsequent authors. The chance of ever locating the Teysmann sheet from Bangka, which therefore must be the type of Mappa cissifolia, seems remote unless a label in Zollinger's hand turns up.
2. This sheet is not in my opinion a Macaranga.
