Malesian Euphorbiaceae Descriptions

67. MACARANGA

 

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)

 

click picture to find more information about Whitmore's book.

 

References

    Davies, S.J. 2001. Systematics of Macaranga sect. Pachystemon and Pruinosae (Euphorbiaceae). Harvard Pap. Bot. 6: 371–448.

    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: 11–87.

    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: 389–394.

    Whitmore, T.C. 2008. The genus Macaranga, a Prodromus. Royal Botanic Gardens Kew.

 

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References

Genus description

Infrageneric and Informal groups:

    Angustifolia group

    Bicolor group

    Brunneo-floccosa group

    Conifera group

    Denticulata group

    Dioica group (plus New Guinean Pseudorottlera)

    Gracilis group

    Javanica group

    Longistipulata group

    Mappa group

    Section Pachystemon

    Section Pruinosae

    Section Pseudorottlera

    Tanarius group

    Section Winklerianae

Regional Keys to the species

    Key to the species the Malay Peninsula

    Key to the species of Sumatra

    Key to the species of Java and Bali

    Keys to the species of Borneo

    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

Species diagnoses

Inadequately known Macaranga names

Excluded Species

 

Macaranga Thouars

 

    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 2–4-locular (3- or 4-locular in Malesia), opening by splits, pistillode absent; in pistillate flowers ovary single, with 1–5 locules, each with 1 ovule, style(s) short or long, usually terminal, occasionally lateral or basal to ovary, stigmas often plumose. Fruits 1–5-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.

 

1. Macaranga acerifolia Airy Shaw (Dioica group)

 

    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 2–4 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 1–3 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 1–3 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.

 

2. Macaranga advena Pax & K.Hoffm. (Gracilis Group)

 

    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 5–6 m. Twigs slender, ferruginous tomentose, becoming glabrous. Stipules ovate, c. 8 mm long, apex acuminate, glabrous, caducous. Leaves: petioles 2–3 cm long, slender, pilose; blades elliptic, 9–20 by 1.5–4 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 6–8 pairs, veins and veinlets reticulate. Staminate inflorescences and flowers unknown. Pistillate inflorescences racemes (? spikes), 5–10 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, 2–3cm long, densely plumose. Fruits and seeds unknown.

    Distribution — Papua New Guinea (Felsspitze).

    Habitat & Ecology — Forest clearings. Altitude: 1400–1500m.

    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.

 

3. Macaranga aenigmatica Whitmore (Group uncertain)

 

    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 20–36 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 1–7 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, 15–20 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.

 

4. Macaranga aλtheadenia Airy Shaw (Section Pachystemon)

 

    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.

 

Macaaeth-habit.gif (117532 bytes)

 

Tree to 15(–25) m tall, 15(–30) cm dbh; twigs 7–10 mm in diam., glabrous, densely glaucous, hollow, housing ants. Stipules ovate, 8–10 by 8–13 mm, recurved, succulent, producing food-bodies on the abaxial surface, 1–3 (rarely more) pairs present on the shoots. Leaves: petioles terete, slender, 15–30 cm long, glau­cous, glabrous; blades broadly to very broadly ovate, large, variable in size, 15–40 by 13–35 cm, usually shallowly trilobed, rarely with just three cusps or with an additional pair of tiny basal leaf cusps, dissected to Ό–½ of the leaf length, central lobe c. 7–12 cm wide with an acuminate apex, lateral lobes ascending with acuminate apices, 2–5 cm peltate, base broadly to very broadly rounded, margin entire, with huge (2–3 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, 14–24 by 9–15 cm, mostly glabrous. distal axes finely ferrugineus pubescent, to 4 axis orders, main unbranched axis flattened and 3–6 cm long, first pair of secondary branches ±opposite with accessory branches; bracts elliptic, 5–8 by c. 5 mm, margin entire, apex acute, glabrous, caducous; flower clusters with c. 15–20 flowers, decussately arranged and strongly grouped at distal ends of inflorescence branches; bracteoles ovate to broadly ovate, c. 4 by 4–5 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 10–1.5 mm long, shortly pedicellate, pedicel not persisting; sepals fused, splitting to c. 1/2, apex glabrous; stamens 1–3; anthers 3-locular. Pistillate inflorescences paniculate, erect, 7–14 by 4–10 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 4–5-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 5–7 by 8–11 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. 3.5–4.0 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 1–2 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 1–3 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.

 

5. Macaranga albescens Perry (Brunneo-floccosa group)

 

    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 10–30 m high. Twigs stout, 3–4 mm diam., strongly angular, coarsely striate, pubescent. Stipules linear-elliptic, c. 4 mm long, pubescent, very soon caducous. Leaves: petioles 3–9 cm long, slender, 1–2 mm diam., terete, pubescent, kneed at both ends; blades ovate, 6–13 by 4–10 cm, leathery, drying grey brown above, pale brown often with whitish patches below, base 5–10 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 10–15 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 4–6, anthers 4-locular. Infructescences open sparsely branched pubescent racemes to 13 cm long; peduncle 1–3 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 2–10 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: 2100—2800 m.

    Note — Stipules described from Perry (1953). Staminate bracteoles variable. Leaves similar to M. trichanthera except for the basal lobe.

 

6. Macaranga aleuritoides F.Muell. (Longistipulata group)

 

    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, 7–11 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 14–32 cm, usually stout, usually strongly D-shaped in transverse section near base and 4–11 mm across, at first tomentose, soon glabrous, drying constricted at base; blades large, strongly trilobed (rarely shallowly lobed or entire ovate), 22–32 by 20–27 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 5–15, anthers 3-locular. Infructescences 15(–26) cm pubescent racemes with sparse short 2 cm distal branches (rarely spicate). Fruits in groups of 1–3 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, 5–7 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.

 

7. Macaranga allorobinsonii Whitmore (Angustifolia group)

 

    Macaranga allorobinsonii Whitmore, Gen. Macaranga Prodr. (2008) 72 — Type: Robinson 1722 (holo K; iso L), Indonesia, Moluccas, Ambon.

 

Tree to 20–30 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 7–11 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 5–6 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 2–3, unilocular, or rarely bilobed, round, 4 mm, wall thin, closely granular glandular and sometimes with a few small warts, style near the base 1–2 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.

 

8. Macaranga amentifera (Brunneo-floccosa group)

 

    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, 3Ί38’S, 143Ί1’E.

 

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, 16–23 by 10–16 cm, thinly leathery, base broadly rounded to truncate, 1–2 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, 5–7 nerves at petiole insertion, tertiaries and quaternaries scalariform. Staminate inflorescences amongst leaves, sparsely branched 5–10 cm panicles, main axis angular 1 mm diam., striate, weakly tomentose or glabrous, peduncle 15 mm, secondary axes short, 1–2 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.

 

9. Macaranga amissa Airy Shaw (Conifera group)

 

    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, 8–14 by 5–10 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 3–4 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 2–3 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.

 

10. Macaranga amplifolia (Mappa group)

 

    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, 17–35 by 9–20 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 3–4 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.

 

11. Macaranga anceps Airy Shaw (Section Pseudorottlera)

 

    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, 9–15 by 3–5.5 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 0–5 mm; bracteoles triangular, 0.5 by 0.4 mm, apex acuminate, spreading, entire, glabrous, persistent, smaller than cluster. Staminate flowers 2–3 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.

 

12. Macaranga andamanica Kurz (Section Pseudorottlera)

 

    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 1.5–3 cm, very slender, terete, glabrous or with pale crisped and spreading hairs; blades elliptic, 7–11 by 2–5cm, 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, 1–1.5 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 3–6 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 1–5 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.

 

13. Macaranga angulata S.J.Davies (Section Pachystemon)

 

    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 4–8 mm in diam., glabrous or rarely with sparse minute hairs, hollow, housing ants, not glaucous. Stipules deltate, 6–10 by 6–10 mm, recurved, succulent, producing food-bodies on abaxial surface, up to 4 pairs present on the shoots. Leaves: petioles terete, 8–20 cm long, glabrous; blades broadly to very broadly ovate, 14–30 by 11–22 cm, deeply trilobed, sometimes with two additional basal leaf lobes or cusps, lateral lobes narrow and spreading, dissected to 1/2–2/3 of the leaf length, 1–3 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 8–9 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, 11–13 by 7–10 cm, glabrous, 3–4 axis orders, main axis flattened, first pair of secondary branches opposite with accessory branches; bracts deltate, 6–10 by 3–4 mm, caducous, margin entire, apex acute, glabrous or with scattered hairs; flower clusters with 20–30 flowers, spirally arranged and evenly spaced along inflorescence branches; bracteoles ovate, 3–5 by 3–4 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, 4–10 by 4–8 cm, glabrous or rarely with minute scattered hairs, up to 2 axis orders; bracts caducous. Pistillate flowers 2.5–3 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 4–5-carpellate; styles c. 5 mm long, fused at base, free and spreading from 1/2–3/4 of the length, persisting to form a prominent 3–4 mm long crown at fruit apex; stigma not dissected. Fruits 5–6 by 9–11 mm, subglobose, sessile, one discrete glandular patch on each carpel wall developing into a slender horn-like process 4–6 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 3–5 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.

  

14. Macaranga angustifolia Lauterb. & K.Schum. (Angustifolia group)

 

    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, 2–3 mm diam., slightly angled and ridged, glabrous, blackish brown. Stipules elliptic, 8 mm, apex acuminate, blackish brown, papery, glabrous, erect, soon caducous. Leaves: petioles 1.2–2 cm, slender, 1–1.5 mm diam., slightly flattened, usually somewhat channelled above, glabrous, weakly kneed at top; blades elliptic, 16 by 3–6cm, 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, 9–13 cm, glabrous or weakly furfuraceous racemes; branches to 4 cm, ascending, some arising from a flower cluster; peduncle 1–3 cm, main axis slender, 1 mm diam., flattened, lower branches alternate or (sub)opposite, thread-like with evenly 2–3 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 1–2 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.

 

15. Macaranga ashtonii S.J.Davies (Section Pachystemon)

 

    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 10–20 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–)15–25 by (8–)10–20 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, 1–3 pairs present on the shoots. Leaves: petioles terete but drying slightly flattened, stout, (12–)15–35 cm long, 4–8 mm in diam., greyish, densely pubescent with erect silvery hairs; blades broadly ovate, large to very large, 25–60 by 15–40 cm, trilobed, dissected to c. 1/4–1/2 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, 4–9 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–)8–9 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, 5–11 by 3–6 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, 1–6 by 3–4 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, 10–15 by 6–12 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 4–6 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 4–5-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 2–3 mm long crown at fruit apex; stigma not dissected. Fruits (immature) 10–12 by 6–8 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 3–6 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, 3–4-thecate anthers, 4–5-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.

 

16. Macaranga astrolabica Pax & K.Hoffm. (Dioica group)

 

    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 2–4 cm, glabrous; blades elliptic, 8–17 by 3.5–5.5 cm, subcoriaceous, base rounded narrowly cordate, with 1–3 glands near petiole insertion, margin entire or wavy, long acuminate, granular-glandular, glabrous, with 3 basal nerves and 6–7 pairs of secondaries, reticulations visible below. Staminate inflorescences panicles 4–8 cm long, narrow, sparsely branched and flowered; peduncle glabrous; branches puberulous; bracts ovate triangular, aepx acute, puberulous without glands; bracteoles elliptic, 2–3 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.

 

17. Macaranga auctoris Whitmore (Group uncertain)

 

    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, 5–7 by 4–5 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, 50–70 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.

 

18. Macaranga baccaureifolia Airy Shaw (Section Pseudorottlera)

 

    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 8–11 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 swamp–for. S. Johore Singapore, 1978) notes that in Johore this is gregarious and loses its leaves when the annual flooding recedes every February.

 

19. Macaranga balabacensis Pax & K.Hoffm. (Longistipulata group)

 

    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 10–15 mm cordate, or truncate, upper side with c. 6 glands between petiole insertion and blade. Staminate bracteoles 5–7 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).

 

20. Macaranga bancana (Miq.) Mόll.Arg. (Section Pachystemon)

 

    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.

 

Macabanc-habit.gif (106860 bytes)

 

Abundant small tree to 23 m tall, but reproductive from only 2–3 m tall; twigs 4–9 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, 6–8 by 6–9 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 2–4 pairs present on the shoots. Leaves: petioles terete, slender, 8–24 cm long, not glaucous, glabrous; blades broadly ovate, 16–32 by 12–25 cm, trilobed to rarely just tricusped, variably dissected to 1/8–1/2 of the leaf length, central lobe broad sometimes constricted near the base, lateral lobes ascending and 2.5–5(–7) cm wide, rather shallowly peltate, petiole insertion 1–4 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 7–8 prominent veins, 2° venation scalariform and rather closely spaced usually with 14–16 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, 10–20 by 5–15 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, 3–6 by 2–3 mm, margin entire, apex acute, finely ferrugineus pubes cent on both surfaces and the margin, caducous; flower clusters with 15–25(–30) flowers, spirally arranged and scattered along ultimate inflorescence branches: bracteoles small, broadly ovate to transversely elliptical, 1.5–2.5 by 2–4 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, 3–11 by 3–7 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, 5–8 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, 1.0–1.5 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 2–3 mm long crown at fruit apex; stigma not dissected. Fruits 5–7 by 10–11 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. 1–2 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.

 

21. Macaranga barkeriana Whitmore (Longistipulata group)

 

    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 10–15m. Twigs very coarsely densely hispid with spreading pale golden 2–3 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 17–36 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 1540–1590m.

    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.

 

22. Macaranga beccariana Merr. (Section Pachystemon)

 

    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.

 

macabecc-habit.gif (124113 bytes)

 

Tree, 15–17 m tall, with simple architecture of the main trunk and secondary branches only; twigs 7–8 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, 9–12 by 2.5–4.5 mm, coriaceous, spreading, not producing food-bodies, all pairs usually persistent on the shoots after leaf fall. Leaves: petioles terete, slender, 10–30 cm long, shorter towards shoot apex, glabrous, glaucous, strongly double pulvinate; blades ovate, 13–30 by 12–20 cm, very deeply and narrowly trilobed, dissected almost to the petiole insertion or 4/5–7/8 of the leaf length, leaf lobes elliptic, 2.5–4 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, 2–4 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 8–10 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, 7–20 by 5–10 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, 3–6 by 3–4 mm, enclosing the flower clusters, margin finely evenly pectinate with 10–16 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, 5–14 by 2–5 cm, slender, up to 2 axis orders, secondary branches ± opposite, covered in minute ferrugineus scaly hairs; bracts elliptic, c. 5–7 mm long, entire, covered in ferrugineus hairs, caducous. Pistillate flowers c. 2–3 mm long, solitary in bracteole axils; calyx urceolate, c. 1.5 mm long, minutely ferrugineus hairy, persistent; ovary 3-carpellate, 1–2 mm long; styles c. 1 mm long, free, spreading, persistent; stigma not dissected. Fruits subglobose, 4.5–6 by 5–6 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 5–11 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.

 

23. Macaranga belensis L.M.Perry (Longistipulata group)

 

    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 1.1–2.3 cm, soon caducous. Leaves: petioles 1–3.5 cm; blades ovate, 5–13 by 2–5.5cm, narrowing to a retuse or emarginate base with 2–6 glands near petiole insertion, tapering gradually to a long 1–2.5cm 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 5–7.5 cm spikes; peduncle 1–2.5 cm; flower clusters spaced; bracteoles ovate, long-acuminate, pilose outside, without patellar glands inside. Staminate flowers in clusters of 7–10, pilose; stamens 11–24, 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.

 

24. Macaranga bicolor Mόll.Arg. (Bicolor 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 5–17 cm, slender, terete, velvety; blades ovate, 8–18(–26) by 7–13(–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 4–12 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.

 

25. Macaranga bifoveata J.J.Sm. (Dioica group)

 

    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 4–18 cm, slender, terete, shallowly striate, sparsely pubescent, kneed at top; blades almost as broad as long, ovate 12–21 by 10–23 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, 4–7 cm long; peduncle 3–6 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 1–3 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 3–5, anthers 4-locular. Pistillate inflorescences spikes or sparsely branched racemes to 7 cm, main axis 2 mm diam., terete, weakly striate, sparsely tomentose; peduncle 2–3 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 1–3, 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 2–3 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).

 

26. Macaranga brachytricha Airy Shaw (Tanarius group)

 

    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, 20–25 by 12–15 mm, sides straight, apex acute, with faint parallel nerves, warm brown, papery, at first weakly scurfy, becoming glabrous, recurved, persistent. Leaves: petioles 9–24 cm, to 5 mm diam. near base, terete, faintly ridged, soon becoming glabrous, strongly constricted at base, weakly kneed at top; blades ovate, 19–30 by 11–24 cm, papery, base broadly (shallowly) rounded, 4–6 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?) 1–3 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.

 

27. Macaranga brevipetiolata Airy Shaw (Section Pseudorottlera)

 

    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.

 

Macabrev-habit.gif (64300 bytes)

 

Treelet or tree 2–10(–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, 2–3 cm, terete, densely pubescent, kneed at both ends; blades obovate, 8–18 by 4–7 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 3–4 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 3–4 cm. Pistillate flowers 3–7. 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.)

 

28. Macaranga brunnea-floccosa Pax & K.Hoffm. (Brunneo-floccosa group)

 

    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, 10–20(–33)m tall, or rarely a 3 m shrub. Twigs slender, 2–3 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 1–4 cm, slender, 1 mm diam., terete, at first rufous floccose, kneed at both ends; blades ovate to ovate-elliptic, 4–9(–13) by 2.5–5.5(–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 3–4 axis orders, usually rufous floccose, branches divaricate or ascending; peduncle 2–10 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 1–3 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, 800–2100m.

    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.

 

29. Macaranga caladiifolia Becc. (Section Pachystemon)

 

    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, 1–3 m tall; twigs 6–12 mm in diam., glabrous, prominently swollen and constricted at some but not all nodes, hollow, housing ants. Stipules narrowly ovate, 8–13 by 3–5 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, 8–22 cm long, glabrous; blades narrowly ovate, 15–28 by 7–14 cm, entire or sometimes with small lateral cusps, 4–6 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 5–7 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, 5–8 by 3–4 mm, persistent, margin entire, apex acute, glabrous; flower clusters with 1–5 flowers, spirally arranged and crowded at the apex of the inflorescence; bracteoles ovate, 3–5 by 2–4 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 1–2 mm long, pedicellate; sepals free, very thin, glabrous; stamens (3–)4–5; anthers 4-locular. Pistillate inflorescences erect, unbranched, with the flowers clustered at the distal end of the rhachis, 2–5 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. 2–3 mm long, fused at base, free and spreading from c. 1/3 of the length, persisting to form a prominent 2–4 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 4–5 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 1–2 m in height, but M. havilandii has broadly ovate recurved stipules, unistaminate flowers, and 4–5 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 15–20 m height and is largely restricted to peat swamp forests, furthermore it has globose evenly glandular 3–4-carpellate fruits, and pubescent inflorescences.

 

30. Macaranga calcicola Airy Shaw (Section Pachystemon)

 

    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 4–10 m tall with glossy green leaves; twigs 4–7 mm in diam., often drying whitish, covered in long erect silvery hairs, hollow, housing ants. Bark smooth, greyish brown. Stipules ovate, 6–8 by 7–9 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, 8–20 cm long, finely pubescent particularly towards the base and apex; blades, very broadly triangular-ovate, 12–24 by 8–20 cm, tricusped to shallowly trilobed, dissected to 1/5 of the leaf length, central lobe triangular 6–15 cm wide at the base and narrowing gradually towards the apex, lateral lobes c. 1–3 by 1–3 cm at the base shortly acute and spreading, 1.5–4 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 7–8 prominent veins, 2° venation scalariform with 9–11 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, 5–15 by 3–8 cm, basal unbranched rhachis very short and only 0.5–3.5 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, 2–3 axis orders, main axis flattened, first pair of secondary branches ±opposite with or without accessory branches; bracts elliptic, 5–7 by c. 3 mm, margin entire, apex acute, finely pubescent particularly towards the base and on the margins, caducous; flower clusters dense with 20–50 flowers, spirally arranged and clustered towards the ends of ultimate inflorescence branches; bracteoles deltate, 3–6 by 2–3 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, 1.5–6 by c. 1.5–3.5 cm, usually with one pair of short opposite branches only, appearing trifid, bracts caducous, not seen. Pistillate flowers 5–7 by 2.5–3.0 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, 3–4 mm in diam.; styles 3–4 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 3–4 mm long crown at fruit apex; stigma smooth. Fruits subglobose, 6–7 by c. 7–10 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).

 

31. Macaranga carrii L.M.Perry (Brunneo-floccosa group)

 

    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 6–24m. Twigs stout, 4–6 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 3–8 cm, diam. 1.5–4 mm, angled, striate, variously orange-brown tomentose, kneed at top; blades ovate to deltoid, size variable, 7–16 by 5–14 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 2–4 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 2–5 cm, peduncle 2–4 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 4–10 mm peltate, apex acute only rarely caudate, tomentum below sometimes dark chocolate brown. Staminate inflorescences with big 5–10 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, 2250–2900 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 20–28m. Stipules carinate especially towards tip. Leaves: blades with base (4–)7–10(–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, 2200–2880m.

    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.

 

32. Macaranga caudata Pax & K.Hoffm. (Longistipulata group)

 

    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 5–9 mm base, golden-tomentose, papery, erect, subpersistent, sometimes splitting at tip. Leaves: petioles 6–15 cm, slende, 2 mm diam., terete, tomentose, weakly kneed at top; blades orbicular deltoid, 12–23 by 10–20 cm, papery, base broadly cordate to truncate, cordulate at petiole insertion, with few-numerous small glands, margins with small prominent glandular teeth, apex 2–7 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.

 

33. Macaranga celebica Koord. (Group uncertain)

 

    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 10–15 cm, fairly slender, round, glabrous, kneed at top; blades ovate elliptic, broadest near base, 16–24 by 5–11cm, 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, 8–10 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.

 

34. Macaranga chlorolepis Airy Shaw (Section Pseudorottlera)

 

    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 3.6–20 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 2.5–4 cm, slender, 1.5 mm diam., terete, very finely scurfy lepidote, kneed at apex; blades ovate-elliptic, 10–19 by 4.5–6cm, 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 17–22, anthers 4-locular. Pistillate inflorescences 2–4 cm spikes, with 2–3 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 30–600m.

    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.

 

35. Macaranga choiseuliana Airy Shaw (Mappa group)

 

    Macaranga choiseuliana Airy Shaw, Kew Bull. 32 (1978) 412; Whitmore in Airy Shaw, Kew Bull., Addit. Ser. 8 (1980)136–8 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, 30–50 by 20–40 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, 8–15 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, 1–2 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 2–5 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.

 

36. Macaranga chrysotricha Lauterb. & K.Schum. (Longistipulata group)

 

    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 5–10 cm, terete, 3 mm diam., variously golden tomentose, sometimes weakly kneed at top; blades deltoid, 13–17(–36) by 9–14(–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 2–4 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.

 

37. Macaranga clavata Warb. (Tanarius group)

 

    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 1–2 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, 8–27 by 6–24 cm, papery, base broadly rounded, 2–7 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 6–9 cm; main axis bearing 2 cm spaced fascicles of up to 5, 1–6 cm long, variously divaricate branches; peduncle 1–3 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, 3–5 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 1–2(–3) lobed, lobes round, 5 mm across, thinly woody, bearing 2–4 mm long blackish spreading rigid viscid conspicuously clavate hairs, swollen towards apex; pedicel slender length variable to 3 cm; styles subulate, 2–3 mm, caducous. Seeds round, 5–6 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.

 

38. Macaranga clemensiae L.M.Perry (Brunneo-floccosa group)

 

    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.

 

Macaclem-habit.gif (69386 bytes)    Macaclem-male.gif (32458 bytes)    Macaclem-fruit.gif (30824 bytes)

 

Trees 11–15 m. Twigs stout, 3–4 mm diam., angled, densely tomentose or pubescent. Stipules elliptic to ovate, 7–15 by 3–5 mm, apex acute, thick, densely dark brown pubescent, soon caducous. Leaves: petioles 4–6 cm, stout, 2 mm diam., terete, densely pubescent, kneed at both ends; blades ovate, 9–15 by 4–8 cm, thickly leathery, base broadly rounded, 2–5 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 4–7 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 1–2 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, 1–2, 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, 1800–2200m.

    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.

 

39. Macaranga coggygria Airy Shaw (Brunneo-floccosa group)

 

    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, 5–7 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, 23–30 by 16–19cm, leathery, base broadly rounded to truncate, 1–3 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 1–3, globose, 3 mm diam., leathery, black granular glandular, with a few short caducous spines; pedicel thread-like, 8–14 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, 1200–1900m.

    Notes — 1. Male inflorescence only known from a fragment.

2. The bracteoles are highly distinctive.

 

40. Macaranga congestiflora Merr. (Bicolor group)

 

    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 4–5 in leaf axils, ultimately to 4 cm; main axis slender, rounded, scurfy, with a few distant, brittle 10–15 mm branches and a few 5 mm tertiary branches, sometimes with axillary branches; bracts subpersistent, ovate 1.5 mm, hispid; flower clusters 3 mm, 3–5 crowded apically; enclosed by persistent, furfurescent, eglandular, papery, nerved bracteoles; latter ovate to oblong, 1.5–3 by 2–7 mm, conspicuous, apex divided to halfway or more into tapering, regular fimbriate strips. Staminate flowers 4–10 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 7–20 mm; sepals small, persistent, free; stigmas persistent, tiny. Seeds 3, spherical, pitted.

    Distribution — Philippines (Palawan).

    Habitat & Ecology — Lowlands.

 

41. Macaranga conifera (Rchb.f. & Zoll.) Mόll.Arg. (Conifera group)

 

    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.

 

macaconi-habit.gif (105332 bytes)    Macaconi-male.gif (38758 bytes)    Macaconi-fruit.gif (83085 bytes)

 

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, 3–4 cm racemes (or with a few short third order axes); main axis terete; peduncle 5–20 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.

 

42. Macaranga constricta Whitmore & Airy Shaw (Section Pachystemon)

 

    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 6–10 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, 5–8 by 5–7 mm, the pair not completely encircling the twig, recurved, succulent, abaxial surface producing food-bodies, 1–3 pairs present on the shoots. Leaves: petioles terete, 10–20 cm long, densely glaucous, glabrous; blades ovate, rather consistent in size and shape, 15–28 by 10–20 cm, shallowly trilobed, dissected to 1/4 to 1/3 of the leaf length, central leaf lobe 5–8 cm wide at the base, lateral lobes ascending 3–6 cm wide, 2–6 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 6–7 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, 18–28 by 12–18 cm, axes mostly glabrous and densely glaucous, distal branches scattered with minute ferrugineus hairs, to 4 axis orders, main axis flattened c. 1/3–1/2 of total inflorescence length, first pair of secondary branches ± opposite with accessory branches; bracts 5–7 by 2–3 mm, persistent, margin entire, apex acute, glabrous or scattered with ferrugineus hairs; flower clusters with c. 20–25 flowers, spirally arranged and clustered on ultimate inflorescence branches; bracteoles ovate-elliptic, 5–7 by 3–4 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 1–1.5 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, 5–14 by 4–10 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, 4–5.5 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 3–5 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 3–4 mm long crown at fruit apex; stigma not dissected. Fruits subglobose, 9–11 by 5–7 mm, pedicellate, with one discrete glandular patch on each carpel wall developing into a slightly raised disc covered in yellow, sticky exudate, glaucous; pedicel 4–6 mm long, glabrous or with scattered minute ferrugineus hairs, glaucous. Seeds subtriangular-ovoid, 4–4.5 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.

 

43. Macaranga costulata Pax & K.Hoffm. (Javanica group)

 

    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.

 

macacost-habit.gif (104046 bytes)    Macacost-male.gif (45538 bytes)    Macacost-fruit.gif (48441 bytes)

 

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 4–7(–12) cm, papery, base broadly truncate occasionally 1–3 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, 3–10 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, 5–10 mm apart; bracteoles linear, 3–4 mm, distorted by 1–3 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 1–3, 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.

 

44. Macaranga crassistipulosa Pax & K.Hoffm. (Angustifolia group)

 

    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 5–6 mm long, apex acute, persistent, thick, glabrous. Leaves: petioles 3–5 cm; blades oblong or ovate-oblong, 7–12 by 2.5–6 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 6–7 mm [?cm] long, divaricate-racemose, stipe short, glabrous; rachis compressed; bracteoles triangular-semiorbicular, 1–2 mm long, apex obtuse or acute, glabrous, 5–9-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.

 

45. Macaranga cucullata J.J.Sm. (Longistipulata 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 3–3.5 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).

 

46. Macaranga cumingii Mόll.Arg. (Javanica group)

 

    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 2–7 cm, slender terete, finely furfuraceous to glabrous; blades ovate to elliptic, 8–12 by 3–6 cm, papery, base cuneate to truncate, sometimes with 1–2 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, 5–8 cm spikes or occasionally with a few short branches; axes slender, furfuraceous; flower clusters small, evenly spaced; bracteoles inconspicuous, size and shape variable, ovate, 1–3(–7) mm long, base sometimes narrowed, apex acute or (long) acuminate, with 2–4 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, 6–13 cm, axes slender, furfuraceous becoming glabrous; flower clusters 1–2 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 2–3(–5), globose, 4–5 mm, smooth, thinly woody, at first with a waxy coating, sometimes with warty excrescence; pedicel thread-like, very long, 15–30 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 — 1000–2200m.

    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 1908–1910, 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.

 

47. Macaranga darbyshirei Airy Shaw (Tanarius group)

 

    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, 4–8(–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 9–17(–20) cm, slightly flattened, 2–4 mm diam., striate, (strongly) glaucous, glabrous or weakly pubescent; blades ovate to ovate oblong, 15–20(–40) by 8–14(–37) cm, leathery, base rounded (sometimes deeply so, with blade widest at middle), deeply 3–5(–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, 3–4 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 4–6 stamens with anthers 4-locular. Infructescences short, 6–11 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, 5–10 mm; calyx caducous. Seeds globose, 6 mm, coarsely irregularly rugose, with sarcotesta.

    Distribution – New Guinea.

    Habitat & Ecology — Lowlands to 900m, primary or secondary forest.

 

48. Macaranga daviesii Takeuchi (Longistipulata group) (W. Takeuchi)

 

    Macaranga daviesii Takeuchi, Harvard Pap. Bot. cxc (2007) 389, fig. 1–4. — 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, 15–20 m tall; bole straight, erect, unbuttressed; crown numerously branched. Wood straw, dense. Twigs terete, subapical diam. (2–)2.5–3.5(–4) mm, firm, not pithy, resinous, exudate black, vernicose, obscurely striate, elenticellate; internodes (0.5–)1–4(–5.5) cm. Indumentum acicular hairs, 2–3 mm long, appressed (or patent), easily dislodged with handling, irritant, dense, persisting, orange-brown (fresh pink on all vegetative parts); scale glands 0.1–0.2 mm diam., hyaline-yellow or pellucid. Stipules united, apparently single, sheathing, (32–)40–62(–76) by 6–10 mm, terminal, erect, caducous, indument as twigs. Leaves: petioles (7–)15–35(–42) by 1–3 mm, planoconvex, pulvinate at both ends or not, black, appressed hairy; blades ovate-rhomboid, (4.5–)7–13(15.5) by (2.5–)6–10.5(12.5) cm, rugulose, coriaceous, base broadly rounded, notched 2–4 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 5–8 pairs above a basal digitate nervation, straight-divergins 25–40Ί 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–)24–36(–40) by 0.8–1.2 mm, black, strigose, granulose-glandular; bracts 4 in 2 dimorphic series, contiguous with calyces; major (outer) bracts: stipitate, trullate, (7–)10–13 by 3–5 mm inclusive a 2–4 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 2–3 mm, persisting in fruit, hirtellous; calyx globose-ovoid, 4–5 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 3.5–4 mm, scurfy, laxly hirtellous, bilocular; styles 2, (5–)7–9 mm long, arcuate, plumosely lobed. Fruits inserted on accrescent peduncles to 65 by 2 mm, globose or ovoid, 9–13 by 10–14 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.

 

49. Macaranga decipiens L.M.Perry (Dioica group)

 

    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 2–3 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, 2–3 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 2–3, anthers 4-locular. Pistillate flowers calyx not seen; ovary 2-locular, style 3–5 mm, plumose. Infructescences spikes, 3–13 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 1–2 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.

 

50. Macaranga densiflora Warb. (Dioica group)

 

    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.62A–E; 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 1.5–16 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', 4–9 cm long; branches ascending to divaricate, thread-like, to 3 cm long; peduncle 2.5–4 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 3–8 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 4–5 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 3–15 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 1–2 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.

 

51. Macaranga denticulata (Blume) Mόll.Arg. (Denticulata group)

 

    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, 3–6(–10) by 1–2(–3) mm, densely fawn furfuraceous, leathery, erect, very soon caducous. Leaves: petioles 5–11 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 10–15 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.

 

52. Macaranga depressa (Mόll.Arg.) Mόll.Arg. (Section Pachystemon)

 

    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.

 

macadepr-habit.gif (120655 bytes)

 

Small tree to 10 m tall, but often smaller or shrub-like; twigs slender, 3–7(–8) mm in diam., solid, not housing ants. Bark smooth, pale grey-brown. Stipules deltate, 6–12 by 5–11 mm, the pair not completely encircling the twig, leathery, not succulent, spreading and usually perpendicular to the twig, usually 4–8 or more pairs present on the shoots. Leaves: petioles terete, slender, 8–20 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, 5–14 by 4–7 mm, margin entire, apex acute, caducous; flower clusters with 20–40 flowers, spirally arranged and grouped towards the ends of inflorescence branches, mostly much larger than the bracteoles; bracteoles small, broadly ovate, 3–4 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, 4–12 by 4–6 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 3–5 mm long, solitary in bract axils; calyx urceolate, 1–2 mm long, densely ferrugineus pubescent, with scattered yellow granular glands towards the apex, persistent, splitting irregularly as ovary expands; ovary 4–5-carpellate, c. 1.5 mm long; styles 2–3 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 3–5 mm long crown at fruit apex; stigma not dissected. Fruits subglobose, compressed, 6–8 by 8–11 mm, sessile, one discrete glandular patch on each carpel wall developing into a long slender horn-like process 3–7 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, 10–35 by 9–32 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 1–3.5(–4) cm, lateral lobes spreading widely and sharply, leaf lobes narrow 1–6 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 7–9 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, 10–35 by 9–30 cm, deeply 3- to 5-lobed, dissected to 4/5 of the leaf length, usually shallowly peltate to 1–4(–5) cm, lateral lobes narrow and spreading widely, leaf lobes narrow 3–6 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–)8–10 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, 5–14 by 4–7 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 2–3 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, 10–30(–35) by 8–25 cm, variably trilobed, sometimes almost entire to shortly tricusped to quite deeply trilobed and sometimes with additional basal cusps or lobes, peltate to (1–)2–6 cm, lateral lobes varying from short acute cusps to spreading narrow lobes 2–5 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 7–9 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.

 

53. Macaranga didymocarpa Whitmore (Conifera group)

 

    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 2–3 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 3–4 mm axis with strongly overlapping bracteoles; latter oblong, 1–2 mm long, broadly acute, sometimes coarsely toothed, papery, pubescent, erect, persistent, longer than and enclosing a tight cluster of 3–4 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.

 

54. Macaranga diepenhorstii (Miq.) Mόll.Arg. (Conifera group)

 

    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, 13–23(–30) by 10–18(–23) cm, papery, base broadly rounded, 2–4(–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, 2–5 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 4–8 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 2–3 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.

 

55. Macaranga dioica (Forst.) Mόll.Arg. (Dioica group)

 

    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.

 

Macadioi-habit.gif (307666 bytes)

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 3–4 mm, acuminate, papery, tomentose, erect, subpersistent. Leaves: petioles 7–13 cm, 2–3 mm diam., terete, tomentose; blades ovate to deltoid, 7–21 by 6–20 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, 2–10 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 2–4 minor nerves at petiole insertion. Staminate inflorescences narrow tomentose racemes to 11 cm; branches divaricate to 15 mm long, 1–2 cm apart, arising from a flower cluster; peduncle 3–4 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.

 

56. Macaranga domatiosa Airy Shaw (Gracilis Group)

 

    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.

 

macadoma-habit.gif (112304 bytes)

 

Treelets or small trees, 3–8(–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 5–6 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 2–3-flowered irregularly spaced clusters. Staminate flowers reddish, developing in sequence, large, sessile; sepals 2, red-nerved, 2–3 mm long, acuminate; stamens 12–25, anthers 4-locular. Pistillate inflorescences 5–15 cm delicate spikes, with 3–4 subulate 1–2 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 4–5 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, 2060–3000m.

    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.

 

57. Macaranga ducis Whitmore (Dioica group)

 

    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 5–25 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, 12–15 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, 13–17 by 8–11cm, thinly leathery, base broadly rounded to truncate to broadly shallowly cordate, with 1–several 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, 4–8 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 2–3, subtended by caducous leafy elliptic 15 mm bracts. Fruits round, to 1 cm diam., leathery, with dense soft flat almost glabrous spines; pedicel stout 1–3 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.

 

58. Macaranga endertii Whitmore (Javanica group)

 

    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.

 

Macaende-habit.gif (49362 bytes)

 

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 6–11 cm, slender, terete, finely pubescent or glabrous; blades broadly ovate, 8–16 by 5–12 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, 6–12 cm long, narrow open racemes with 10–25 mm distant slender divaricate side branches, without axillary branches, finely pubescent or glabrous; main axis flattened; peduncle 15–40 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, 2–3 mm long, bearing 1–few 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.

 

59. Macaranga eymae L.M.Perry (Longistipulata group)

 

    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, 14–26 by 4–7 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 1–5 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 6–7 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).

 

60. Macaranga fallacina Pax & K.Hoffm. (Longistipulata group)

 

    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, 12–22 by 5 mm, rusty pubescent, papery, caducous. Leaves: petioles 8–12 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, 1–1.5 cm peltate with 3–5 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, 3–4 axis orders, furfuraceous or glabrous; main axis flattened, 1.5 mm diam.; peduncle 1–3 cm; some branches opposite, without axillary branches, sometimes arising from a flower cluster, secondaries divaricate to ascending, 4–10 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 4–5 cm distant clusters of 1–2 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.

 

61. Macaranga fragrans L.M.Perry (Mappa group)

 

    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 2–4 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 1–2 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 34–43 cm panicles, peduncle 5–12 cm. Fruits 2(3)-locular, tomentellose, loculi 9 mm long by 6 mm, with 0.5–2 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).

 

62. Macaranga galorei Whitmore (Dioica group)

 

    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 4–8 cm, terete, 1.5–3 mm diam., densely pale tomentose; blades deltoid, broader near base 12–22 by 8–17 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, 5–13 cm spikes, axis terete to slightly flattened, pubescent, with an apical and sometimes 1–2 subapical clusters of 3–6 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, 3–8 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.

 

63. Macaranga gigantea (Rchb.f. & Zoll.) Mόll.Arg. (Section Pruinosae)

 

    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.

 

macagiga-habit.gif (122802 bytes)    Macagiga-male.gif (45530 bytes)    Macagiga-fruit.gif (63032 bytes)

 

Enormously abundant pioneer tree to 30 m tall and 30–40 cm dbh, usually smaller and flowering from 5–10 m; twigs stout, 10–25 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, 15–60 by 10–40 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 20–40(–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, 20–60 by 20–50 cm, often much larger in saplings, thickly coriaceous, trilobed, dissected to 1/3–1/2, deeply peltate to 2–13 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 5–8 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, 15–50 by 10–30 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, 4–10 by 2–5 mm, apex acute, finely silvery pubescent on both surfaces, persistent; flower clusters with 6–15 flowers, spirally arranged and usually in groups of 2–6 on the ultimate inflorescence branches; bracteoles ovate to broadly ovate, 4–6 by 2–5 mm, margin deeply irregularly dentate with 4–8 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 2–3; anthers 4-locular. Pistillate inflorescences paniculate, erect, 12–25 by 6–12 cm, densely branched, densely silvery pubescent on all axes, 3 axis orders; bracts ovate, 3–5 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 4–5 by 6–8 mm, bilocular, locules subglobose, pedicellate, evenly covered in yellow sticky glandular exudate, without processes, with scattered silvery hairs between the locules; pedicel 4–6 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. 800–1000 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 10–20 years following the abandonment of shifting cultivation (Kochummen, Malayan Forester 29, 1966, 170–181). 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.

  

64. Macaranga glaberrima (Hassk.)Airy Shaw (Section Pseudorottlera)

 

    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, 1.5–3 mm diam., terete, glabrous. Stipules and bud scales subulate, 5 mm, chestnut brown, glabrous, erect, crowded at twig tip, soon caducous. Leaves: petioles 2.5–3 cm, slender, 1 mm diam., terete, glabrous, with a small conspicuous apical knee; blades elliptic, 8–10(–15) by 2.5–4(–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 450–1100 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).

 

65. Macaranga glandibracteolata S.J.Davies (Section Pachystemon)

 

    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.

 

macaglan-habit.gif (122345 bytes)

 

Tree to 25 m tall and c. 35 cm dbh, with an open spreading crown; twigs 5–12 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, 6–8 by 5–7 mm, producing food-bodies on abaxial surface, 1–3 pairs present on shoots. Leaves: petioles terete, slender, 18–25 cm long, scattered to evenly covered with minute silvery hairs particularly towards the apex; blades broadly ovate, 25–45 by 16–35 cm, >3 cm peltate, shallowly tricusped to trilobed, dissected to 1/10–1/5 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 7–8 prominent veins, 2°–4° venation prominent and densely scalariform; young leaves finely red-brown pubescent. Staminate inflorescences paniculate, erect, 20–30 by 20–30 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, 4–7 mm long, ± persistent, margin entire, apex acute, rarely terminated by a small conical nectary, abaxial surface densely pubescent; flower clusters of 4–10 flowers decussately arranged in groups of 6–10 at branch apices; bracteoles ovate, 3–6 by 3–4 mm, enclosing the flowers, margin finely pectinate with the central lobe elongated 1–2 mm with a small conical nectary at apex, apex acute, abaxial surface finely pubescent. Staminate flowers 1–1.5 mm long, sessile; sepals splitting to the base, apex pubescent; stamens 3–4; anthers 3-locular. Pistillate inflorescences paniculate, erect, 5–12 by 5–10 cm, stout, up to 3 orders of branching, secondary branches ± opposite, very finely red-brown pubescent; bracts caducous. Pistillate flowers 3.5–5.5 by 2–3 mm, solitary in bracteole axils, pedicel developing with the fruit; calyx c. 4 mm long, urceolate, densely and finely red-brown pubescent, persistent; ovary 2–3 by c. 2 mm, 4–5-carpellate, minutely pubescent at the apex and on the sutures; styles 1–2 mm long, fused at the base, free and spreading from 1/4–1/3 of the length, persisting to form a 2 mm long crown on the fruit apex; stigma not dissected. Fruits subglobose, 5–7 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 4–8 mm long, glabrous or scattered with minute ferrugineus hairs, glaucous. Seeds subtriangular-ovoid, 4–5 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, 3–4-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.

 

66. Macaranga gracilis Pax & K.Hoffm. (Gracilis Group)

 

    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 3–10 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, 4.5–9.5 by 1.5–2.5 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 2–10 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 1–3 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, 1500–2200 m. Rare.

    Note — I place Sayer 12528 here. It differs in 4–5 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.

 

67. Macaranga grandifolia (Blanco) Merr. (Mappa group)

 

    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 2–5 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 4–8 per cluster, shortly pedicellate, 0.8 mm diam.; sepals free;, stamens 2–3(–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 Hawai’i (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.

 

68. Macaranga griffithiana Mόll.Arg. (Section Pachystemon)

 

    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 4–5 m tall; twigs 4–9 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, 4–6 by 6–7 mm, the pair not completely encircling the twig, recurved, succulent, producing food-bodies on the abaxial surface, 1–4 pairs present on the shoots but often caducous. Leaves: petioles terete, slender, 6–15(–23) cm long, glaucous, glabrous; blades ovate, 10–25(–28) by (8–)10–23 cm, usually shallowly trilobed, dissected to 1/5–1/3 of the leaf length, central lobe broadly deltate c. 4–8(–10) cm wide at the base, lateral lobes 1–3(–6) cm wide short and spreading with shallow rounded sinuses, 2–5 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 6–7(–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, 10–25 by 10–18 cm, basal axes mostly glabrous and densely glaucous, distal axes sparsely to densely ferrugineus pubescent, up to 4 axis orders, main unbranched axis 2–8 cm long flattened, first pair of secondary branches ± opposite with accessory branches; bracts elliptic, 7–15 by 3–7 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, 3–5 by 2.5–4.0 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 2–3; anthers 4-locular. Pistillate inflorescences paniculate, erect, 4–16 by 3–8 cm, stout, basal axes glabrous and densely glaucous, distal axes covered in fine ferrugineus hairs, 2–3 axis orders, secondary branches ± opposite without accessory branches; bracts caducous, not seen. Pistillate flowers c. 3–4 mm long, solitary in bract axils; calyx urceolate, 2–3 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 2–3 mm long crown at the fruit apex; stigma not dissected. Fruits subglobose, compressed, 7–9 by 8–12 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 2–4 mm long, scattered with minute ferrugineus hairs, glaucous. Seeds subtriangular-ovoid, 3.5–4 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 4–5-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.

 

69 Macaranga hartleyana Whitmore (Angustifolia group)

 

    Macaranga hartleyana Whitmore, Gen. Macaranga Prodr. (2008) 144. — Type: Hartley 10694 (holo K), Papua New Guinea, Central Prov., ‘Western Section’, Kakoda Road at Owen’s 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 2–4 cm, slender, 1 mm diam., terete, striate, densely finely pubescent, weakly kneed at both ends; blades elliptic, 6.5–12 by 2–4 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 1–3 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, 660–1000m.

 

70. Macaranga havilandii Airy Shaw (Section Pachystemon)

 

    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 3–5(–6) m tall and 2.5 cm dbh with no or few secondary branches and leaves clustered at the apex of the stem; twigs 5–8 mm in diam., glabrous, usually densely glaucous, hollow, housing ants. Stipules ovate, 3–6 by 4.0–6.5 mm, not completely encircling the twig, recurved, succulent, producing food-bodies on abaxial surface, 2–6 pairs present on the shoot. Leaves: petioles terete, very long and slender, 12–40 cm long, glabrous; blades ovate, 20–35 by 13–22 cm, shape very variable, usually deeply trilobed or 5-lobed with broad basal cusps or rarely sub-entire, usually dissected to 1/2–3/4 of the leaf length, deeply peltate to 5–10 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 6–7 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, 7–12 by 2–4 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, 8–12 by 4–6 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, 5–7 by 2–3 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, 7–16 by 2–8 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 4–6 mm long, solitary in bracteole axils; calyx urceolate, 1.5–2 mm long, with scattered fine ferrugineus hairs, persistent; ovary 4–5-carpellate, c. 1.5 mm long; styles 3–5 mm long, fused for 4/5 of length, free and spreading at apex, persisting to form a prominent 3–5 mm long crown at fruit apex; stigma not dissected. Fruits subglobose, 5–6 by 8–11 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 3–7 mm long, slender, glaucous, glabrous or with fine scattered ferrugineus hairs. Seeds 4–4.5 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 3–5 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 3–5 stamens, 4-locular anthers, stems that are irregularly swollen, and is not glaucous.

 

71. Macaranga hengkyana Whitmore (Brunneo-floccosa group)

 

    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.

 

72. Macaranga herculis Whitmore (Tanarius group)

 

    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.

 

73. Macaranga heynei I.M.Johnson  (Javanica group)

 

    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 5–8 cm, slender, 1 mm diam., terete, finely ridged, weakly rusty furfuraceous or glabrous, weakly kneed; blades trullate, 10–18 by 4.5–8 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.

 

74.  Macaranga hispida (Blume) Mόll.Arg. (Longstipulata group)

 

    Macaranga hispida (Blume) Mόll.Arg. in DC., Prodr. 15, 2 (1866) 990; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 366, t. 61A–E; 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.

 

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Small trees, 4–10 m tall with irritant hairs. Twigs tetragonal or flattish near tip, stout, 4–7 mm across, finely warm brown hispid, rarely (sub)glabrous. Stipules united, apparently single, erect, leathery, big and prominent, elliptic, 5–9 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, 15–3–(–40) by 7–20(–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 7–15 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 3–6 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, 8–14(–26) cm long, axis flattened, 3 mm wide; peduncle 7–15 cm; bracts ovate elliptic, 15 by 3–8 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, 10–15 mm diam., leathery, shaggy with long fine tapering hirsute soft processes, surface granular glandular; pedicel stout 2–3 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.

 

75. Macaranga hoffmannii L.M.Perry (Dioica group)

 

    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, 10–20 m tall. Twigs slender, dark, glabrous and sparsely granular-glandular. Stipules not seen, most certainly small or minute. Leaves: petioles 2–3.5 cm, slender, glabrous; blades ovate or ovate-elliptic, 7–12 by 2.5–4 cm, leathery, base narrowed and narrowly peltate, 3-nerved base plus 2–4 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 4–6 cm panicles, divaricately branched almost at base, glabrous, granular-glandular, branches slender of more or less equal length; clusters 10–20 flowered; lower bracteoles oblong, 2 mm long, upper triangular, denticulate. Staminate flowers 2 mm broadl sepals 3, elliptic, acute, outside densely granular-glandular; stamens 10–12, anthers 4-locular. Pistillate unknown.

    Distribution — Northeast New Guinea (known from the syntypes only).

    Habitat & Ecology — Montane forests, 1400–1500 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.

 

76. Macaranga hosei King ex Hook.f. (Section Pruinosae)

 

    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.

 

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Large tree with a bluish-green appearance, to 20–30 m tall and 30–40 cm dbh, although flowering somewhat smaller; twigs 6–11 mm in diam., terete, glabrous, usually densely glaucous. Bark pale grey, white or light brown, prominently hoop-marked, smooth or minutely fissured. Stipules 10–15 by 10–15 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, 10–25 cm long, slightly glaucous, glabrous; blades broader than long to almost rounded, not flat, slightly convex, 8–20 by 10–25 cm, deeply trilobed, dissected to c. 1/2–3/4, the lateral lobes almost the same size as the central lobe, the lateral lobes slightly asymmetrical, not peltate or rarely 1–5 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 2–3 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, 20–40 by 10–20 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, 5–7 mm long, margin entire, glabrous, often persistent; flower clusters with 6–15 flowers, spirally arranged on inflorescence branches; bracteoles broadly ovate, 4–6 by 3–4 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 2–3; anthers 4-locular. Pistillate inflorescences erect narrow panicles often extending above the leaves, 15–35 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, 5–10 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, 4–5 by 5–6 mm, green or yellowish-green at maturity, locules subglobose, without processes, covered in yellowish-green, sticky exudate; pedicel slender, 5–8 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.

 

77. Macaranga hullettii King ex Hook.f. (Section Pachystemon)

 

    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.

 

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Small tree to 18 m tall, but often flowering from as small as 3 m; twigs 5–9 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, 5–9 by 5–10 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. 3–4 pairs persistent on the stem. Leaves: petioles terete, 8–20(–25) cm long, mostly glabrous or sometimes finely silvery pubescent; blades narrowly ovate to ovate, 14–32 by 6–16(–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, 1–3(–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 7–9 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, 10–30 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 1/2–3/4 of total inflorescence length, first pair of branches opposite with accessory branches; bracts ovate-elliptic, 6–13 by 3–6 min, margin entire, apex acute, glabrous or scattered with fine ferrugineus hairs towards the base, caducous; bracteoles ovate-elliptic, 4–8 by 3–5 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 20–35 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 4–10 by 2–5 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 3–5 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 2–3 mm long, expanding at maturity, fused at the base, free and spreading from c. 2/3 of the length, persisting to form a prominent 3–5 mm long crown at fruit apex. Fruits subglobose, 6–8 by 10–13 mm, sessile, one discrete glandular patch on each carpel wall developing into a long slender horn-like process 4–7 mm long, covered in greenish-yellow, sticky exudate. Seeds ovoid, 4–4.5 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.

 

78. Macaranga hypoleuca (Rchb.f. & Zoll.) Mόll.Arg. (Section Pachystemon)

 

    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.

 

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Tree to 30 m tall and 30–40 cm dbh, flowering somewhat smaller; twigs 7–12(–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, 10–30 cm long, glabrous, glaucous; blades broader than long to almost rounded, usually slightly convex, 10–24 by 12–30 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 1.5–4 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 8–10 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 10–28(–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, 3–6 mm long, margin entire, both surfaces scattered with minute ferrugineus hairs; bracteoles ovate, 2.5–4 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 10–15 per cluster, c. 1 mm long, shortly pedicellate; sepals 3, fused, densely covered in minute ferrugineus hairs; stamens 1; anthers 4-locular. Pistillate inflorescences 5–15 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, 5–7 mm long, entire or with a finely pectinate apex, covered in ferrugineus hairs, caducous. Pistillate flowers c. 2–3 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, 5–6 by 6–8 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 3–11 mm long, glaucous, with scattered minute crisped ferrugineus hairs. Seeds 3–4 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. 

 

79. Macaranga hystrichogyne Airy Shaw (Brunneo-floccosa group)

 

    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, 4–6 m. Twigs 3 mm diam. near tip, terete, densely persistently pale brown tomentose. Stipules triangular, 3 by 3–7 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, 14–20 by 14–20 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 6–10, 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, 2–4 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, 450–1230 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.

 

80. Macaranga indica Wight (Denticulata group)

 

    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, 7–11, 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 6–18 cm, slender, to 3 mm diam., terete, smooth, glabrous, not kneed; blades ovate, 10–22 by 9–20 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, 6–12 cm long, usually 2 occasionally 3 axis orders; peduncle 2–4 cm; axes (strongly) flattened, to 2 mm wide, glabrous or furfuraceous, striate; primary branches 1–3 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 1–2 mm stalk and a big conspicuous subapical round 2–3 mm diam. patellar gland, occasionally twin glands; flower clusters often on a 1–2 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 5–8, 0.5 mm diam., shortly pedicellate; sepals free, glabrous except often with a few subapical granular glands; stamens 5–7, 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 5–10 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).

 

81. Macaranga indistincta Whitmore (Section Pachystemon)

 

    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.

 

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Small tree to 15 m tall with an overall rather dull green appearance; twigs 5–9 mm in diam., densely pubescent with fine erect silver-grey hairs, hollow, housing ants. Bark pale brown, smooth. Stipules ovate, 6–9 by 8–10 mm, dark brownish to purple when fresh, recurved, succulent, adaxial surface finely pubescent, producing food-bodies on abaxial surface, 2–6 pairs present on shoots. Leaves: petioles terete, 10–22 cm long, densely pubescent; blades ovate to broadly ovate, 15–32 by 14–25 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, >2–3 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 7–9 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, 9–27 by 6–18 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, 4–7 by 3–4 mm, margin entire, apex acute to acuminate, glabrous, caducous; flower clusters with 15–25 flowers, spirally arranged and evenly spaced along inflorescence branches; bracteoles broadly ovate, 2–3.5 by 2.5–4 mm, cupulate and enclosing flower clusters, glabrous, margin pectinate with 6–12 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, 6–15(–20) by 5–10 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 3–4 mm long, solitary in bracteole axils; calyx urceolate, 2–2.5 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, 2–3 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 2–3 mm long crown at the fruit apex; stigma not dissected. Fruits subglobose, 4–6 by 7–10 mm, sessile, one discrete glandular patch on each carpel wall developing into a long slender horn-like process 4–7 mm long, covered in yellowish green, sticky exudate. Seeds 3.5–4 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.

 

82. Macaranga induta L.M.Perry (Brunneo-floccosa group)

 

    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 10–17 m. Twigs 3 mm, squarish, densely pubescent to furfuraceous, becoming subglabrous. Stipules oblong linear, 3–6 mm, apex acute, very soon caducous. Leaves: petioles 2–5 cm, 1.5 mm diam., terete, striate, at first finely pubescent, kneed at both ends; blades elliptic to ovate-elliptic, 7–17 by 3–7cm, broadest near base, leathery, base cuneate to rounded, 2–4 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 8–14 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 6–7 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 2–4 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, 1300–2300 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 4–20 m. Blades very soon glabrous below and only 5–8 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, 1400–2600 m.

 

83. Macaranga inermis Pax & K.Hoffm. (Angustifolia group)

 

    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 3–4 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 3–15 cm long, 1.5–3 mm diam., terete, glabrous or weakly furfuraceous, kneed at both ends but more strongly at top; blades ovate, less often elliptic, (10–)15–20(–26) by (4–)9–14(–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 1–2 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 1–4, 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, 3–8 mm; calyx persistent; style basal, papillose, caducous, 3–8 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, 0–2000 m. Common.

    Note — A widespread and polymorphic species.

 

84. Macaranga intonsa Whitmore (Brunneo-floccosa group)

 

    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 25–36 by 19–34 cm, leathery, base broadly shallowly cordate or 5 cm peltate, margin sinuate and minutely distantly denticulate, apex abruptly caudate, 3–6 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, 1200–2100 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.

 

85. Macaranga involucrata (Roxb.) Baill. (Dioica group)

 

    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. 62F–G; 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, 2–10 m. Twigs slender, 3 mm diam., terete, finely ochreous pubescent, or occasionally (sub)glabrous. Stipules setose or narrowly elliptic, 8–12 by 1(–2) mm, ochreous tomentose, erect, caducous. Leaves: petioles 3–8 cm, slender, terete, usually densely (rarely sparsely) ochreous pubescent, some hairs tufted; blades broadly ovate, 10–16 by 8–12 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 5–10 cm, dense to open racemes, with distant divaricate often slender branches, distally sometimes thread-like; axes terete, densely finely pubescent; basal peduncle short, 2–4 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, 8–18-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 1–5 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 3–10, anthers 4-locular. Pistillate inflorescences pendulous 5–10 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, 10–15 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.

 

86. Macaranga javanica (Blume) Mόll.Arg. (Javanica group)

 

    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 5–7 cm, slender, 1.5 mm diam., terete, weakly furfuraceous; blades ovate oblong, 9–13 by 4–7 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 1–3, weakly bilobed, 3 by 4 mm, black granular glandular and with sparse tiny blunt caducous spines, thinly woody; pedicel 3–4 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.

 

87. Macaranga johannium Whitmore (Mappa group)

 

    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.

 

88. Macaranga kinabaluensis Airy Shaw (Javanica 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 5–27 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 5–9 cm, slender, terete, very soon glabrous, not kneed; blades ovate or trullate-ovate, 7–15 by 4–9(–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 1–4 adaxial patellar glands; some persisting, flower clusters 5–15 mm apart, subtended by bracteoles; latter linear to subspatulate, 5(–10) by 1 mm, furfurescent, with 1–4 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, 1–3 mm, with 1 or a few patellar glands and finely sparsely furfuraceous, mostly caducous. Pistillate flowers c. 1 cm, apart in clusters of 1–3; pedicel 1–3 mm; calyx with 2–3 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, 200–1700 m, sometimes on ultrabasics.

 

89. Macaranga kingii Hook.f. (Section Pachystemon)

 

    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 7–13 mm diameter, glabrous, moderately glaucous, hollow, housing ants. Stipules narrowly ovate, 15–26 by 8–13 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, 15–40 cm long, usually slightly glaucous, glabrous; blades ovate to broadly ovate, 20–55 by 20–40 cm, adaxial surface dark shiny green, appearing slightly bullate, very deeply dissected with 5–7 widely spreading lobes, deeply peltate to 4–10 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 7–8 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, 10–20 by 8–12 mm, persistent, margin entire, apex acute, glabrous; flower clusters with 1–5 flowers, spirally arranged and crowded towards the ends of inflorescence branches; bracteoles ovate, 4–6 by 3–4 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 4–7; anthers 4-locular. Pistillate inflorescences paniculate, erect, to 10 cm long, stout, glabrous, usually glaucous, 1–2 axis orders, secondary branches ± opposite; bracts caducous, not seen. Pistillate flowers c. 12 mm long, solitary in bract axils; calyx urceolate, 4–6 mm long, not glaucous, with scattered ferrugineus hairs near the base, persistent, splitting irregularly as the ovary expands; ovary 4–5(–6)-carpellate, 4–6 mm long; styles 4–9 mm long; styles fused at base, free and spreading from c. 1/2 of the length, persisting to form a prominent 6–8 mm long crown at the fruit apex; stigma shallowly dissected. Fruits subglobose, 9–12 by 10–15 mm, pale pink to bright red, pedicellate, two discrete glandular patches on each carpel wall developing into narrow ridges raised to 1–2 mm, covered in dark, sticky exudate; pedicel 10–30 mm long, often finely pubescent, not glaucous. Seeds 5.5–6.5 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 5–7 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.

 

90. Macaranga kostermansii L.M.Perry (Gracilis Group)

 

    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, 2.5–4.5 by 1–1.2 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 0–3 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, 1600–2300 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.

 

91. Macaranga laciniata Whitmore & Airy Shaw (Javanica group)

 

    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 2–3 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.

 

92. Macaranga lamellata Whitmore (Section Pachystemon)

 

    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.

 

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Small tree to 15 m tall, flowering from c. 3 m; twigs 8–15 mm in diam., glabrous, moderately to densely glaucous, hollow, housing ants. Stipules narrowly ovate, 15–27 by 8–13 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, 15–50 cm long, up to 8–10 mm in diam., usually glaucous, glabrous; blades ovate, often huge, 20–65 by 14–36 cm, entire or rarely with very short lateral cusps, 4–10 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 7–9 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, 4–12 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, 8–15 by 5–10 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, 4–6 by 3–4 mm, enclosing flower clusters, margin ± entire or rarely with 1–2 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 4–5; anthers 4-locular. Pistillate inflorescences paniculate, erect, 5–14 by 3–6 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 4–5(–6)-carpellate; styles fused at base, free and spreading from c. 3/4 of the length, persisting to form a prominent 6–10 mm long crown at fruit apex; stigma not dissected. Fruits subglobose, 9–12 by 10–18 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. 2–4 mm, covered in yellow, sticky exudate; pedicel 5–18 mm long, glabrous, usually glaucous. Seeds subtriangular-ovoid, 5.5–6.5 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.

 

93. Macaranga lanceolata Pax & K.Hoffm. (Angustifolia group)

 

    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 2–3 cm, slender, 1 mm diam., terete, glabrous or with a few spreading hairs, kneed at top; blades (narrowly) oblong elliptic 5–12 by 1.5–3.5 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 4–5, sessile; stamens 4, anthers 4-locular. Pistillate inflorescences as staminate. Pistillate flowers solitary; pedicel 3–5 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, 850–1500 m.

    Note — Like M. faiketo the leaves and fruits lack granular glands and the latter are furfuraceous. The leaf base is distinctive.

 

94. Macaranga leytensis Merr. (Mappa group)

 

    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).

 

95. Macaranga lineata Airy Shaw (Tanarius group)

 

    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 15–20; 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.

 

96. Macaranga loheri Elmer (Javanica group)

 

    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 4–5 cm, slender, terete, glabrous; blades ovate, 7–14 by 4–7 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.

 

97. Macaranga longicaudata L.M.Perry (Brunneo-floccosa group)

 

    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, 7–10 by 4–7 cm, leathery, base truncate sometimes very broadly very shallowly cordate, usually with 4–10 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, 4–6 by 4–5 cm plus basal peduncle of 3–6 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 8–9, globular, 1 mm, granular glandular; stamens 4, anthers 4-locular. Fruits with slightly eccentric single style.

    Distribution — East New Guinea.

    Habita & Ecology — Mountain forests, 2400–2800 m.

    Note — Similar to M. induta but easily told apart by the leaf base and strong bullation.

 

98. Macaranga longistipulata (Kurz ex Teijsm. & Binn.) Mόll.Arg. (Longistipulata group)

 

    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, 5–12 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, 15–24 by 4–6 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, 1000–1500 m.

    Note — A big montane tree, highly distinctive in its stipule, leaf nervation and fruit ornamentation.

 

99. Macaranga louisiadum Airy Shaw (Dioica group)

 

    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, 2–3 mm diam., slightly angled, finely pubescent. Stipules subulate, 6 mm, erect or somewhat recurved, pubescent, subpersistent. Leaves: petioles 2–3 cm, slender, 1–2 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 1–2 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 1–5 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.

 

100. Macaranga lowii King ex Hook.f. (Section Pseudorottlera)

 

    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 53/7–11, 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.

 

Macalowi-habit.gif (95967 bytes)    Macalowk-habit.gif (77723 bytes)

 

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 3–7 cm, slender, 1 mm, terete, weakly pubescent or glabrous, kneed; blades elliptic, 8–15 by 3–5 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 3–4 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 1/3–2/3 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, 2–4 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!

 

101. Macaranga lugubris Whitmore (Dioica group)

 

    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.

 

102. Macaranga lumiensis Whitmore (Gracilis Group)

 

    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 2–3 mm apart, evenly spaced, each compound cluster of 2–3 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.

 

103. Macaranga magnifolia L.M.Perry (Mappa group)

 

    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, 2–10 cm long, 0.7–3 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, 24–67 by 16–60 cm, papery or occasionally leathery, base broadly rounded, 4–17 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 2–6 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 3–4 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.

 

104. Macaranga mappa (L.) Mόll.Arg. (Mappa group)

 

    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 7–20 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 2–3 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.

 

105. Macaranga melanosticta Airy Shaw (Brunneo-floccosa group)

 

    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, 6–20 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, 7–12 by 4–6 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 8–10 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 3–4 mm, sessile; sepals papery granular glandular; stamens 9, anthers 4-locular. Infructescences spikes to 6 cm; peduncle to 4 cm, terete, granular glandular. Fruits 1–4 per infructescence, globose, to 6 mm diam., leathery, finely pubescent and black granular glandular; pedicel stout. eventually 1 cm; calyx persistent; styles 2, apical, 3–5 mm, persistent, smooth. Seeds slightly rugulose, with sarcotesta.

    Distribution — New Guinea.

    Habitat & Ecology — Secondary or primary montane forest, 2400–3300 m.

 

106. Macaranga minahassae Whitmore (Tanarius group)

 

    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, 20–50 by 15–20 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, 6–12 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, 3–5 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 6–7, 0.3–0.5 mm; pedicel 0.3–0.5(–1) mm; stamens 4–7; 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.

 

107. Macaranga misimae Airy Shaw (Gracilis Group)

 

    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, 2–3 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 5–10 mm, slender, 1 mm diam., terete, densely tomentose; blades elliptic to oblong-elliptic, 8–9.5 by 2–3 cm, thinly leathery, tapering gradually to a finely cordate base with 2–7 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 11–13 pairs, well-spaced, looping and joining near margin, lower part of lamina often somewhat bullate. Staminate inflorescences small delicate sparsely branched 3–7 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 6–8 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, 2–3 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).

 

108. Macaranga motleyana (Mόll.Arg.) Mόll.Arg. (Section Pachystemon)

 

    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].

 

Macamotl-habit.gif (112419 bytes)    macamotl-male.gif (37266 bytes)    macamotl-fruit.gif (46426 bytes)

 

Slender glaucous tree to 20 m tall with branches clustered towards the ends of the axes, flowering from 4–5 m tall in open areas; twigs 6–9(–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, 4–7 by 6–7 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, 2–4 pairs present on the shoots but often caducous. Leaves: petioles terete, slender, 10–20 cm long, glaucous, glabrous or sometimes with scattered minute ferrugineus hairs; blades ovate, (15–)20–35 by (10–)12–25 cm, usually deeply trilobed, sometimes with an additional pair of basal leaf cusps or lobes, dissected to 1/3–1/2 of the leaf length rarely less, central lobe c. 4–8 cm wide usually quite narrow near the base, often with ± parallel sides, lateral lobes c. 3–7 cm wide ascending and remaining quite close to the main lobe with narrow sinuses, 2–5(–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 6–7(–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, 10–22(–25) by 10–18 cm, basal axes mostly glabrous and densely glaucous, distal axes sparsely to densely ferrugineus pubescent, up to 4 axis orders, main unbranched axis 2–8 cm long flattened, first pair of secondary branches ± opposite with accessory branches; bracts elliptic, 7–17 by 3–6 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, 3–5 by 2.5–4 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 2–3; anthers 4-locular. Pistillate inflorescences paniculate, erect, 4–10 by 3–5(–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. 5–7 by 4–5 mm, solitary in bract axils; calyx urceolate, 3–4 mm long, densely covered with fine ferrugineus hairs and scattered yellow granular glands, persistent, splitting irregularly as ovary expands; ovary 4–5(–6)-carpellate, c. 3 mm long; styles c. 2–3 mm long, fused at base, free and spreading from c. 1/2 of the length, persisting to form a prominent 3–5 mm long crown at the fruit apex; stigma not dissected. Fruits subglobose, compressed, 7–9 by 9–13 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 4–8 mm long, scattered with minute ferrugineus hairs, glaucous. Seeds subtriangular-ovoid, 3.5–4 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.

 

109. Macaranga myriantha Mόll.Arg. (Angustifolia group)

 

    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, 10–16 by 3–4.5 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 7–12 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 1–4 short flat spines; pedicel 4–10 mm, slender; sepals elliptic, 1 mm, persistent; style basal, breaking off. Seeds unknown.

    Distribution — Moluccas (Morotai, Buru, Ambon).

    Habitat & Ecology — Forest, 900–1000 m.

 

110. Macaranga neobritannica Airy Shaw (Dioica group)

 

    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, 10–13 by 3–5 mm, apex acuminate, chestnut brown, papery, at first both sides tomentose, erect, subpersistent. Leaves: petioles 4–10 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, 15–23 by 8–13 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, 2–3 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 2–4 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 1–3, 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.

 

111. Macaranga noblei Elmer (Mappa group)

 

    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, 16–30 by 15–28 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 2–4 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, 4–5 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 2–4 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.

 

112. Macaranga novoguineensis J.J.Sm. (Dioica group)

 

    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, 2–3 mm diam., weakly angled, densely usually flavous pubescent. Stipules (narrowly) elliptic, 10 by 1 mm, slightly curved, pubescent, erect, subpersistent. Leaves: petioles 3–6 cm, slender to fairly stout, 2–4 mm diam., terete, pubescence as twigs, kneed at top; blades ovate to broadly ovate, 12–16 by 7–12 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 15–20 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 5–10 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, 500–990 m.

    Note — This lower montane, glabrous, narrower leaved variety is distinctive. The only staminate specimen seen does not have the flower clusters crowded.

 

113. Macaranga nusatenggarensis Whitmore (Tanarius group)

 

    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 8–10 cm, slender, round, striate, golden furfuraceous to subglabrous; blades round to deltoid, 11–16 by 7–11 cm, papery, base broadly rounded or less commonly abruptly truncate, 2–3 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 5–6(–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 1–3 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, 1–3 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.

 

114. Macaranga ovatifolia Merr. (Mappa group)

 

    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).

 

115. Macaranga pachyphylla Mόll.Arg. (Bicolor group)

 

    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 G–DC; 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 8–17 cm, slender, 2 mm diam., terete, weakly to densely tomentose with at least a few spreading hairs; blades ovate, 9–18(–30) by 6–13(–25) cm, coriaceous, base broadly rounded, 2–8 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 1–4 cm; main axis slender, 1.5–3 mm diam., terete or flattened, coarsely ridged; lower branches often opposite, sometimes alternate, sometimes with a few axillary branches; bracts variable, ovate, 3–10 by 1–5 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, 1–4 by 1–3 mm, usually with abrupt 1 mm caudate tip, margin entire, both sides densely rusty furfuraceous. Staminate flowers 4–6, 0.5 mm diam.; pedicel 0.5 mm;, sepals hairy; stamens 3–4, anthers 4-locular. Infructescences 4–10 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–)500–1500 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, 500–1500 m.

    Note — This glabrous or nearly glabrous variety is partially sympatric with the type variety whose habitat it shares.

 

116. Macaranga palustris Whitmore (Brunneo-floccosa group)

 

    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 6–7 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.

 

117. Macaranga papuana (J.J.Sm.) Pax & K.Hoffm. (Longistipulata group)

 

    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 2–3.5 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, 9–21 by 4.5–14 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 4–5, 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.

 

118. Macaranga parabicolor Whitmore (Bicolor group)

 

    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 10–13(–40) cm, round or angled, 3 mm diam., striate, pruinose or rusty furfuraceous; blades very broadly ovate, 18–45 by 17–40 cm, thinly leathery, base very broadly rounded to subtruncate, deeply 5–12 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 2–4 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, 100–1200 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.

 

119. Macaranga pearsonii Merr. (Section Pruinosae)

 

    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.

 

macapear-habit.gif (115962 bytes)

 

Large tree to 35 m tall and 50(–60) cm dbh, often smaller in open areas; twigs 6–12 mm in diam., glabrous, usually densely glaucous, hollow, housing ants. Bark pale whitish-grey to light brown, smooth, hoop marked. Stipules broadly ovate, 9–14 by 9–13 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, 10–30 cm long, glaucous towards the base, finely pubescent towards the apex; blades ovate to very broadly ovate, 13–28 by 13–30 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 1–6 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, 5–9 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, 15–40 by 10–25 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, 5–10 by 3–6 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, 3–6 by 2–4 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 2–3; anthers 4-locular. Pistillate inflorescences paniculate, erect, 8–25 by 8–16 cm, densely branched, finely silvery pubescent throughout, up to 3 axis orders; bracts ovate, 5–10 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 3–4 by 5–6 mm, bilocular, locules globose, pedicellate, evenly covered in sticky glandular exudate, without processes; pedicel 4–7 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.

 

120. Macaranga pentaloba S.J.Davies (Section Pruinosae)

 

    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, 18–30 mm in diam., terete though strongly ribbed when dry, densely covered in 2–3 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 2–3 mm long simple erect hyaline hairs, persistent in a cluster at the shoot apex. Leaves: petioles 30–50 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, 40–70 by 30–60 cm, probably even larger in saplings, thickly coriaceous, 5–7 lobed, dissected to c. 1/3 with ovate to broadly ovate lobes, deeply peltate to 5–12 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 5–7 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.

 

121. Macaranga pepysiana Whitmore (Section Pseudorottlera)

 

    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, 2–3 mm long, densely tomentose, erect, persistent. Leaves: petioles 1 cm, terete, 1 mm diam., glabrous, strongly kneed at both ends; blades elliptic, 10–17.5 by 2.5–4.5 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 2–3, 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, 300–610 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.

 

122. Macaranga petanostyla Airy Shaw (Section Pachystemon)

 

    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, 7–12 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, 8–11 by 10–12 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–)15–30 cm long, not glaucous, glabrous; blades glossy dark green, broadly to very broadly ovate, 20–40(–50) by 15–40 cm, trilobed or tricusped and rarely some specimens with an additional pair of basal cusps, dissected to 1/8–1/4 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, 5–9 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 8–10 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, 6–19 by c. 10 cm, basal unbranched rhachis very short 0.5–3 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, 6–16 by 3–8 mm, margin entire, apex acute, mostly glabrous or scattered with fine ferrugineus hairs towards the base, caducous; flower clusters with 15–20 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, 2–4 by 1.0–1.5 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, 7–12 by 6–10 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 3–4 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 4–5 carpellate, c. 3 mm long; styles 3–5 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 5–8 mm long crown at the fruit apex with the style apices remaining curled; stigma not dissected. Fruits (not quite mature) subglobose, compressed, 6–9 by 10–13 mm, sessile, one discrete glandular patch on each carpel wall developing into a long slender horn-like process 4–6 mm long, covered in glandular, sticky exudate. Seeds subtriangular-ovoid, 4–4.5 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.

 

123. Macaranga pilosula Airy Shaw (Tanarius group)

 

    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 6–12 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 12–16 cm, terete, 3 mm diam., faintly ridged, glabrous with grey patches, or sparsely hairy, sometimes constricted at tip; blades ovate-oblong, 19–28 by 11–25 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 25–30; 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.

 

124. Macaranga pleioneura Airy Shaw (Angustifolia group)

 

    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, 8–10 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, 11–17 by 4–7 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 (12–22 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, 1–1.2 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. 30–40, 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 2–8, tiny, 2 mm, globose, at first densely ochreous granular glandular, sometimes puberulous, with 0–6 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, 900–2400 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).

 

125. Macaranga pleiostemon Pax & K.Hoffm. (Angustifolia group)

 

    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, 2–3 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 1–2 cm, 1.5 mm diam., angled, glabrous, weakly kneed at top; blades elliptic to narrowly elliptic, 9–15(–19) by 2–3(–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. 14–16(–20) pairs. Staminate inflorescences sparsely branched 5 cm glabrous to puberulous racemes with 1–3 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 1–2; 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 30–60, 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–)1000–1800 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.

 

126. Macaranga pleytei L.M.Perry (Longstipulata group)

 

    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 3–6 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 2–3, 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.

 

127. Macaranga polyadenia Pax & K.Hoffm. (Angustifolia group)

 

    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.

 

 macapoly-habit.gif (153040 bytes)

 

Small trees or to 18m tall and 30 cm diam., sometimes with stilt roots or plank buttresses. Twigs slender, 2–3 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 6–11 cm, slender, 1.5 mm diam., terete, blackish, usually glabrous, occasionally rusty furfuraceous, weakly kneed at both ends; blades narrowly to broadly ovate, 9–15(–22) by 4–10(–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 6–8 cm long, without axillary branches; secondaries 1 cm distant, to 3 cm long; tertiaries robust, 5 mm. Fruits usually paired, small, 2–3 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 300–600 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.

 

128. Macaranga praestans Airy Shaw (Section Pseudorottlera)

 

    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, 17–28 by 4–10 cm, thinly leathery, base cuneate to rounded with a pair of tiny 1–2 mm convex auricles and 2 conspicuous raised glands, margin undulate often slightly recurved, apically tapering, sometimes abruptly so, to a 2–4(–6) cm caudate tip, drying warm brown, weakly bullate, glabrous with very sparse usually blackish granular glands below, secondary nerves 12–15 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 3–4 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 2–3 cm diam. clusters, spaced along twigs amongst and behind leaves, spikes to 2 cm, axis stout, angled, weakly pubescent and sparsely granular glandular, bearing 1–6 flowers subtended by 2 stipule-like rudimentary bracts; peduncle 2–5 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.

 

129. Macaranga pruinosa (Miq.) Mόll.Arg. (Section Pruinosae)

 

    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].

 

macaprui-habit.gif (124203 bytes)    macaprui-male.gif (47073 bytes)    macaprui-fruit.gif (49021 bytes)

 

Peat swamp tree to 25–30 m tall and 30–40 cm dbh, flowering smaller; twigs 5–12 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, 8–15 by 8–15 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 3–5 pairs persistent on the shoots. Leaves: petioles terete, 10–25 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, 8–25 by 8–27 cm, evenly trilobed with the central lobe slightly larger than laterals, dissected to c. 1/3–1/2, deeply peltate to 1.5–4 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 5–9 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, 6–18 by 4–10 cm, all axes densely minutely silvery or golden ferrugineus pubescent, the basal axis sometimes ± glabrous, 3–4 axis orders, main axis short and terete or slightly flattened, first pair of secondary branches alternate without accessory branches; bracts narrowly ovate, 2–4 by 2–3 mm, margin entire, apex acute, glabrous or scattered with ferrugineus hairs towards the base and on the margins, persistent; flower clusters with 6–15 flowers, spirally arranged on the inflorescence branches; bracteoles ovate to narrowly ovate, 3–5 by 2–3 mm, often with quite straight sides to almost strap-like, margin shallowly irregularly dentate with 4–8 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 2–3; anthers 4-locular. Pistillate inflorescences paniculate, erect, 5–18 by 2–8 cm, sparsely branched, densely ferrugineus pubescent throughout, the hairs slightly less dense on basal axes, 2–3 axis orders; bracts ovate, 3–4 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 5–6 mm, bilocular, locules subglobose, pedicellate, evenly covered in glandular yellow sticky exudate, without processes; pedicel 4–6 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.

 

130. Macaranga puberula Heine (Section Pruinosae)

 

    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 15–20 cm dbh; twigs usually quite stout, 8–15 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, 10–20 by 10–25 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, 15–35 cm long, densely silvery-grey pubescent throughout; blades broadly ovate, 15–30 by 18–35 cm, deeply trilobed, dissected to 1/2–2/3 of the leaf length, lateral lobes spreading 1/2 to 4/5 of the size of the central lobe, 1–3 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 5–7 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, 20–30 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, 4–8 by 3–5 mm, margin entire, apex acute, glabrous or with scattered silvery hairs on the margins and towards the base, persistent; flower clusters with c. 10–15 flowers, spirally arranged on inflorescence branches; bracteoles ovate-elliptic, 2.5–5 by 2–4 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 2–3; anthers 4-locular. Pistillate inflorescences paniculate, erect, 15–35 by 10–20 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 3–4 by 4–5 mm, bilocular, locules globose, pedicellate, evenly covered in sticky glandular exudate, without processes; pedicel 4–7 mm long, densely finely pubescent. Seeds c. 2–2.5 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 800–1800 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.

 

131. Macaranga punctata K.Schum. (Dioica group)

 

    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, 2–3 mm diam., broadly angled, later terete, finely rusty furfurescent, rarely glabrous, dark brown. Stipules (narrowly) oblong-ovate, 3 by 1.5–2 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, 14–23 by 5–13 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, 1–4 cm apart; peduncle 2–5 cm; main axis slender, 1–2 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 2–6 sunken patellar glands, plus a variously developed concave base bearing the flowers, hairy outside. Staminate flowers in clusters of 8–11, maturing sequentially, shortly pedicellate; sepals free, with a few granular glands; stamens 4–7, long-exserted at maturity, anthers 4-locular. Infructescences short spikes or less often racemes to 7(–10) cm, axis flattened, furfuraceous; peduncle 1.5–6 cm; bracts elliptic, 5 mm, leaf-like, rusty pubescent, soon caducous. Fruits solitary, widespaced along axis, plus a terminal group of 2–4, 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.

 

132. Macaranga puncticulata Gage (Bicolor group)

 

    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.

 

macapunc-habit.gif (74026 bytes)    Macapunc-male.gif (50717 bytes)    Macapunc-fruit.gif (33172 bytes)

 

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 6–12 cm, slender, terete, striate, glabrous, glaucous, with a furfuraceous knee at top; blades ovate-oblong, 12–18 by 8–12 cm, thinly coriaceous, base broadly rounded, 2–4 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, 3–8 cm, peduncle 3–4 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, 8–13 by 3–6 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).

 

133. Macaranga quadriglandulosa Warb. (Tanarius group)

 

    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, (10–14(–23) by (6–)12(–18) cm, drying papery and brittle and usually crinkled, base strongly (1–4 cm) peltate, broadly rounded, lower main nerves usually with large flat glands totalling 3–8, 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 4–7 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 4–5(–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.

 

134. Macaranga racemohispida Whitmore (Longistipulata group)

 

    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, 2–3 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.

 

135. Macaranga rarispina Whitmore (Section Pseudorottlera)

 

    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 6–12 cm, slender, 2 mm diam., terete, coarsely striate, glabrous, with a conspicuous apical knee; blades elliptic 16–25 by 6–11 cm, thinly leathery, base rounded or cuneate, with conspicuous gland-bearing auricles 1–5 mm long, margin usually sinuous and with distant small glandular teeth, apex fairly abruptly 10–15 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, 5–8 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 1–2 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.

 

136. Macaranga recurvata Gage (Conifera group)

 

    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.

 

 macarecu-habit.gif (75911 bytes)

 

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 16–20 cm, stout, 3 mm diam. round to angled, striate, glabrous, weakly kneed at both ends; blades ovate-oblong, 22–40(–45) by 16–26(–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 5–10(–20) cm, 4–5 mm wide; main axis stout, flattened, 2–4 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, 1–2 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 5–7, 1 mm diam., pedicel 1 mm; stamens 2–3, 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.

 

137. Macaranga reiteriana Pax & K.Hoffm. (Longistipulata group)

 

    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, 2–3 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, 8–13 by 2–2.8 cm, papery, base rounded with 1–2 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, 1400–2300 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.

 

138. Macaranga rhizinoides (Blume) Mόll.Arg. (Denticulata group)

 

    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 15–40 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 9–12 cm, slender, 2 mm diam., weakly crisped pubescent, constricted near base; blades broadly triangular-ovate to ovate 12–20 by 8–7 cm, leathery, base broadly rounded to truncate, to 1–1.5 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 2–7 mm, contracted at base, lightly dentate, tomentose, caducous; bracteoles fairly narrow triangular-ovate, 2–3 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 7–14, 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).

 

139. Macaranga rorokae Whitmore (Brunneo-floccosa group)

 

    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, 17–25 by 15–23 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 2–3 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 10–15 mm peltate and the reticulations only weakly raised below. It is sterile and conceivably a juvenile.

 

140. Macaranga rostrata Heine (Section Pachystemon)

 

    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, 3–5 mm in diam., terete, glabrous or finely pubescent with short silvery hairs, not glaucous, solid, not housing ants. Stipules ovate-elliptic, 5–10 by 3–5 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, 1–4 pairs present on shoots. Leaves: petioles terete, 3–12 cm long, finely pubescent with short silvery or golden hairs, rarely almost glabrous; blades ovate-elliptic, broadest near the base, 9–20 by 4–10(–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 2–4 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 5–7 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, 4–15 by 1–6 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, 5–12 by 3–6 mm, margin entire, apex acute, mostly glabrous or margin and adaxial surface scattered with minute ferrugineus hairs, caducous; flower clusters with 15–25 flowers, spirally arranged and evenly spaced; bracteoles broadly rounded, 2–4 by 3–5 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, 3–7 by 1–3 cm, stout, 1–2 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 5–6 mm long, solitary; calyx c. 3 mm long, urceolate, glabrous, persistent, splitting irregularly as ovary expands; ovary 4(–5)-carpellate, 3–4 mm long; styles 2–3 mm long, fused at base, free and spreading from c. 1/3 of length, persisting to form a prominent 3–4 mm long crown at fruit apex; stigma not dissected. Fruits subglobose, 6–8 (excluding horns) by 9–10 mm, sessile, one discrete glandular patch on each carpel wall developing into a very long and slender horn-like process 7–15 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 4–5-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.

 

141. Macaranga rufescens S.J.Davies (Section Pruinosae)

 

    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 20–30(–35) m tall and 30–40(–50) cm dbh, although flowering from 3–5 m in open areas; twigs 8–15 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, 12–20 by 12–20 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, 10–40 cm long, slightly glaucous towards the base, glabrous, usually red when fresh; blades broader than long to almost rounded, not flat, slightly convex, 14–25 by 14–35 cm, deeply trilobed, dissected to c. 1/2–3/4, the lateral lobes almost the same size as the central lobe, the lateral lobes slightly asymmetrical, not peltate or rarely 1–5 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 2–3 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, 10–25 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, 5–15 mm long, margin entire, glabrous or rarely with scattered hairs on the margins, often persistent; flower clusters with 6–15 flowers, spirally arranged on inflorescence branches; bracteoles broadly ovate, 4–6 by 3–4 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 2–3; anthers 4-locular. Pistillate inflorescences erect narrow panicles usually not extending above the leaves, 12–28 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, 5–10 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, 4–5 by 5–6 mm, turning red at maturity, locules subglobose, without processes, covered in yellowish-green, sticky exudate; pedicel slender, 5–8 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.

 

142. Macaranga rufibarbis Warb. (Dioica group)

 

    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 2–5.5 cm, slender, 1(–2) mm diam., finely rufous pubescent or glabrous, kneed at top; blades deltoid, size very variable, 10–20 by 7–16 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, 4–15 cm long, rufous tomentose; branches normal to ascending; peduncle 2–3 cm; main axis 1.5–2 mm diam., terete, lowest branches opposite; bracts variable, 3–6 by 1–3 mm, papery, almost glabrous to hirsute, soon caducous; all branches arising from a flower cluster, to 5 cm long; compound flower clusters evenly 5–10 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 1–3 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, 3–10 mm, tomentose; calyx persistent; style apical. Seeds round, coarsely verrucose, with sarcotesta.

    Distribution — Throughout New Guinea

    Habitat & Ecology — Secondary montane forests, 1400–1900 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, 9–17 by 2–6 cm.

    Distribution — East New Guinea (Finisterre Range). Only twice collected.

    Habitat & Ecology — Mountains at 1300 m.

 

143. Macaranga salicifolia Airy Shaw (Longistipulata group)

 

    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 1–5 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 2–3, dissimilar: one spathe-like, 6 by 1–2 mm, apex obtuse, adpressed pilose and granular glandular outside with a terminal gland-bearing limb, plus 1–2 others subulate, 3–5 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.

 

144. Macaranga salomonensis L.M.Perry (Tanarius group)

 

    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, 3–4 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 6–17 cm sparsely branched racemes with 2–7 cm glabrous peduncle; branches 1.5–4 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 5–8, 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.

 

145. Macaranga sandsii Whitmore (Angustifolia 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, 15–18 by 7–8 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, 1–2 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.

 

146. Macaranga sarcocarpa Airy Shaw (Bicolor group)

 

    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 8–18 cm, stout, 3 mm diam., terete, tawny tomentose; blades orbicular-ovate, c. 17 by 14 cm, thickly leathery, base broadly rounded, 4–7 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, 4–10 mm long, furfuraceous; sepals 4 mm, furfuraceous; stigmas 2 mm, apical, persistent, filiform. Seeds globose, 4–5 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.

 

147. Macaranga serratifolia Whitmore (Dioica group)

 

    Macaranga serratifolia Whitmore, Gen. Macaranga Prodr. (2008) 238. — Type: Sands 366 (holo K; iso L), Sulawesi, South, Enrekang, Latimojong Mountains.

 

Small tree. Twigs slender, 1–2 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, 5–12 by 1.5–4 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 1–3 adaxial patellar glands. Staminate flowers 0.3 mm, shortly pedicellate; stamens 2 or 6, anthers 4-locular. Pistillate inflorescences as staminate, spikes 2–5 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, 1100–1650 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.

 

148. Macaranga setosa Gage (Section Pseudorottlera)

 

    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 4–7 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 10–12, anthers 4-locular. Infructescences axis 4–6 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).

 

149. Macaranga similis Pax & K.Hoffm. (Dioica group)

 

    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, 5–10 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, 13–24 by 13–18 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 4–7 cm; main axis fairly slender, 1.5 mm diam., terete, branc