Malesian Euphorbiaceae Descriptions

30. CLEIDION (Euphorbiaceae)

K.K.M. Kulju & P.C. van Welzen

Kulju, K.K.M. & P.C. van Welzen. 2005. Revision of the genus Cleidion (Euphorbiaceae) in Malesia. Blumea 50: 197–219.

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Genus description

Key to the Malesian species

Species description

Doubtful species

Excluded species

Cleidion Blume

    Cleidion Blume, Bijdr. (1825) 612; Müll.Arg., Linnaea 34 (1865) 183; in DC., Prodr. 15, 2 (1866) 983; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 288; in Engl. & Harms. Nat. Pflanzenfam. ed. 2, 19c (1931) 127; Backer & Bakh.f., Fl. Java 1 (1964) 487; Airy Shaw, Kew Bull. 26 (1972) 234; Whitmore, Tree Fl. Malaya 2 (1973) 79; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 74; Kew Bull. Add. Ser. 8 (1980) 57; Kew Bull. 36 (1981) 279; Kew Bull. 37 (1982) 12; Alph. Enum. Euphorb. Philipp. Isl. (1983) 13; McPherson & Tirel, Fl. Nouvelle Caledonie & Depend. 14 (1987) 143; G.L.Webster, Ann. Missouri Bot. Gard. 81 (1994) 86; Govaerts, Frodin & Radcl.-Sm., World Checklist & Bibl. Euphorbiaceae 1 (2000) 356; Radcl.-Sm., Gen. Euphorbiacearum (2001) 211; Kulju & Welzen, Blumea 50 (2005) 200; Welzen & Kulju in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 164; G.L.Webster in Kubitzki, Fam. Gen. Vasc. Pl. 11 (2014) 118, Fig. 27. — Type: Cleidion javanicum Blume.

    Psilostachys Turcz., Bull. Soc. Imp. Nat. Moscou 16 (1843) 58. — Type: Psilostachys axillaris Turcz. [= Cleidion tricoccum (Casar.) Baill., vide Radcl.-Sm. (2001) 212].

    Redia Casar., Nov. Stirp. Bras. (1843) 51. — Type: Redia tricocca Casar. [= Cleidion tricoccum (Casar.) Baill., vide Radcl.-Sm. (2001) 212].

    Lasiostyles C.Presl, Abh. Königl. Böhm. Ges. Wiss. 5, 3 (1845) 579. — Type: Lasiostyles salicifolia C.Presl [= Cleidion javanicum Blume].

    Tetraglossa Bedd., J. Lit. Sci. 2, 22 (1861) 70. — Type: Tetraglossa indica Bedd. [= Cleidion javanicum Blume].

Shrubs to trees; mono- or dioecious; very young branches somewhat angular when dry, older branches terete, glabrescent. Indumentum consisting of simple, short hairs, sericeous (to hirsute with erect or variably oriented hairs), most parts glabrous to only sparsely hairy (the inside of organs always (sub)glabrous); granulate surfaces absent to present. Stipules triangular, usually caducous, base and central part thickened, margin entire and sometimes hairy, apex acute. Leaves alternate, simple; petiole with adaxial groove, usually basally and apically pulvinate, sometimes with 2 raised glands at apex; blade (ovate to) elliptic to obovate, symmetric, papery (to somewhat coriaceous in C. papuanum), margin subentire to crenate to serrate (tooth size varying within a species), with a gland in each tooth, apex usually acuminate; both surfaces with small roundish extra-floral nectaries (= macular glands; in C. moniliflorum only on lower surface), on upper surface mainly grouped loosely near the base, on lower surface mainly in the basal half of the blade and often more or less in lengthwise rows; upper surface usually darker green than lower surface; venation pinnate, ± flat above, raised beneath, nerves looped and closed near the margin, veins (irregularly) scalariform to reticulate, veinlets reticulate. Inflorescences axillary, single (or in C. moniliflorum the staminate ones ramiflorous and often 2 together), unisexual, racemiform (to paniculate outside Malesia) in both sexes to single-flowered when pistillate, rachis often with longitudinal ridges when dry, staminate nodes with 1 flower only or bearing several flowers in a fasciculate to glomerulate (to slightly elongated and branched) cluster, pistillate nodes with 1–3 flowers (or sometimes remaining sterile); bracts triangular to ovate, base and central part thickened, margin entire, often hairy, apex acute (to acuminate when staminate), staminate ones persistent, pistillate ones caducous to persistent; bracteoles resembling bracts but smaller, pistillate ones in pairs at the base of the pedicel (absent when inflorescence single-flowered). Flowers actinomorphic, pedicellate; sepals free; petals and disc absent. Staminate flowers: pedicel with an abscission zone in the middle part; buds broadly ovoid, apiculate; sepals (2 or) 3 (or 4), ovate, valvate, apex acute, mucronulate; receptacle somewhat swollen; stamens 40–120 (in Malesia), filaments flattened, glabrous, anther dorsifixed, 4-celled, on both sides 2 cells above each other, opening introrsely and cruciately, connective often darker in colour, apiculate; pistillode absent. Pistillate flowers: pedicel with abscission zone (absent in C. spiciflorum), widening towards the apex; buds elongated; sepals 3–5, ovate to triangular, imbricate, base and central part thickened, margin entire, often hairy, apex acute, usually persistent; ovary 2- or 3- (or 4-)locular; 1 ovule per locule; style often very short to absent; stigmas deeply split, upper surface papillose. Fruits lobed capsules, opening septicidally and (sometimes incompletely) loculicidally, inner wall usually with shallow depressions; column with a broadened apex. Seeds subglobose, ± smooth, straw to brown when dry, usually mottled, testa not fleshy, caruncle absent.

    Distribution — Pantropical (c. 33 spp.), most diverse in Asia and the South West Pacific. Seven species in Malesia.

Key to the Malesian species

Notes

1. Cleidion javanicum Blume

    Cleidion javanicum Blume, Bijdr. (1825) 613; Kurz,  Forest Fl. Burma 2 (1877) 390; Hook.f., Fl. Brit. Ind. 5 (1885) 444; Brandis, Indian trees (1906) 592; Cooke, Flora Presid. Bombay 2, 3 (1906) 618; Talbot, Forest Fl. Bombay 2 (1911) 488; Pax &  K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 290; Gagnep. in Lecomte, Fl. Indo-Chine 5 (1926) 450; Kanjilal, P.C. Kanjilal, R.N. De & A. Das, Fl. Assam (1940) 220; Backer & Bakh.f., Fl. Java 1 (1964) 487; Prain, Bengal Pl. 2 (1963) 712; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 74; Kew Bull. Add. Ser. 8 (1980) 57; Kew Bull. 36 (1981) 279; Kew Bull. 37 (1982) 12; H.S. Kiu, S.M. Hwang & Y.T. Chang, Fl. Reipubl. Popularis Sin. 44, 2 (1996) 75; Kulju & Welzen, Blumea 50 (2005) 202, fig. 1b, 2, map 1; Welzen & Kulju in Chayam. & Welzen, Fl. Thailand 8, 1 (2005) 167, plate IX: 3; Fl. Thailand 8, 2 (2007) 616, Fig. 7. — Lectotype (designated by Kulju & Welzen, 2005): Blume (Anonymous) s.n. ‘1336’ (holo L, barcode L 0270644), Java, in monte Bonkok Provinciae Tjanjor.

    Lasiostyles salicifolia C.Presl, Abh. Königl. Böhm. Ges. Wiss. 5, 3 (1845) 579. — Type: Helfer 221 (n.v.), Martaban.

    Rottlera urandra Dalzell, Hooker’s J. Bot. Kew Gard. Misc. 3 (1851) 229; Dalzell & A. Gibson, Bombay Fl. (1861) 230. — Type: Dalzell s.n. (K), in jugo Syhadrensi, prope Phondaghat.

    Tetraglossa indica Bedd., J. Lit. Sci. 2, 22 (1861) 70. — Type: Beddome? (n.v.).

    Macaranga tamiana K.Schum., Notizbl. Königl. Bot. Gart. Berlin 2 (1895) 52. —Type: Bammler 4 (n.v.), New Guinea, Kaiser Wilhelmsland, Tami Isl.

    Cleidion spiciflorum auct. non (Burm.f.) Merr. (excl. type): Merr., Interpr. Herb. Amboin. (1917) 322, in obs.; Enum. Philipp. Fl. Pl. 2 (1923) 439; Airy Shaw, Kew Bull. 26 (1972) 234; Whitmore, Tree Fl. Malaya 2 (1973) 79; Airy Shaw, Kew Bull. 35 (1980) 607; Alph. Enum. Euphorb. Philipp. Isl. (1983) 13; Grierson & D.G. Long, Fl. Bhutan 1, 3 (1987) 803; Philcox in Dassan., Revis. Handb. Fl. Ceyl. (1997) 167. See note 1.

(Shrub to) tree, up to 30 m high, dbh up to 50 cm, dioecious (to rarely monoecious); flowering branches 1–4.5 mm thick. Outer bark smooth to slightly roughened or fissured, thin, grey to brown (to greenish), cream to yellow inside, inner bark 3–10(–20) mm thick, light brown to yellowish to straw; sapwood white to straw to creamy yellow. Indumentum sericeous, in staminate parts also hirsute and variably oriented, vegetative parts (excluding domatia) glabrous, generative parts (sub)glabrous to sparsely hairy (except ovary glabrous to densely hairy); granulate surfaces absent. Stipules 2–3.5 by 0.75–1.5 mm, early caducous. Leaves usually in Terminalia growth (flushes); petiole 0.9–9 cm long, usually with 2 raised glands at apex; blade (ovate to) elliptic (to slightly obovate), 5–24 by 1.7–9.7 cm, length/width ratio 1.7–3.8(–4.6), base obtuse to cuneate (to attenuate), margin subentire to serrate with coarse teeth, teeth 7–20 per side, apex (rounded to acute to) acuminate, very apex retuse to rounded to acute, lower surface with shallow, open domatia with single and tufted hairs; nerves (3–)4–10 per side. Staminate inflorescences up to 25 cm long, basally 0.5–1.5 mm thick, flowers 3–several per node, in fascicles, rachis green; bracts 1–2.5 by 1–2 mm. Staminate flowers: pedicels up to 7 mm long, light green; buds 1.5–3 mm diam., brown when dry; sepals 2–4 by 1–2.5 mm, (light) green; stamens 48–120, filaments up to 1.8 mm long, anthers (0.3–)0.4–0.6 by 0.3–0.6(–0.8) mm, yellow to cream, apiculum 0.1–0.3 mm long. Pistillate inflorescences 2.5–11.5 cm long, basally 0.75–1.5 mm thick, 1-flowered, with 1–4 sterile bracts; latter 1.5–2 by 0.75–2 mm, early caducous. Pistillate flowers: abscission zone absent; sepals (4 or) 5, 1–2.5 by 1–2 mm, green, caducous to persistent; locules 2 (or 3); style 1–12 mm long; stigmas erect, 15–30 mm long. Fruits: one locule often reduced (perfect 3-seeded fruits extremely rare), 11–18 by 13–21 (1-seeded) or 19–28 mm (2-seeded), tardily opening, surface ± smooth to verrucose, green when fresh, dark brown to dark grey when dry, inner wall often mottled. Seeds 8.5–14 by 8.5–13.5 mm; hilum elliptic to obovate, 2.5–5.5 by 1–2.5 mm.

    Distribution — From India and Sri Lanka through SE Asia mainland to S China and Indochina, throughout Malesia (absent in the Moluccas) to Australia (Queensland), the Solomon Islands and Vanuatu.

    Habitat & Ecology — In open to shaded places, evergreen to deciduous primary and secondary forests, often along rivers or streams; alluvial to dry soil; varying bedrock, but often on limestone. Altitude up to 1200 m. Flowering and fruiting: Thailand: January to March (fl), April to August (fr); Sumatra & Borneo: March to August (fl & fr); Java: August to December (fl & fr); Philippines: January to April (fl), April to September (fr).

    Vernacular names — Thailand: Di mee (Kariëng); dimi, ma-dee-mee, maj maduug. Sumatra: Urel tenge. Borneo: Entupak (Iban). Java: Hura batu (Sundanese). Bali: Putian (Javanese). Philippines: Paitan, santiki (Tagalog); tubataba (Tagbanua). Papua New Guinea: Tea (Bembi); marramamoi (Kaigulin); la mala goli goli (West Nakanai, New Britain); siwono, ibubu (Bougainville Isl.). Solomon Islands: Saola (Kwara’ae). Vanuatu: Doptop wumer (Uripiv, Malekula Isl.); nagheliromp (Larevet, Malekula Isl.).

    Uses — Various parts are used medicinally in Thailand and the Philippines: a decoction of the poisonous leaves will produce abortion, and a decoction of the bark is used for stomach ache (Quisumbing, Medic. Pl. Philipp., 1978); in the Solomon Islands a decoction of the bark is used externally to treat scabies (Henderson & Hancock, Guide Useful Plants Solomon Isl., 1988). It is a common village shade tree in the Solomon Islands (Henderson & Hancock, Guide Useful Plants Solomon Isl., 1988). The wood is used for house posts in Vanuatu. The wood is reported to be greyish white and soft (Burkill, Dict. Econ. Products .Malay Penins., repr., 1, 1966).

    Notes — 1. The two epithets javanicum and spiciflorum have been inconsistently used for this species after Merrill (1917) interpreted the type of Acalypha spiciflora Burm.f. as belonging to it. This interpretation was based on the figure of Burman’s type in Flora Indica (Burman, Flora Indica, 1768: 203). Later, in his annotation of the actual type in the Delessert Herbarium (G), Merrill identifies the type as a species of Claoxylon and not at all as Cleidion. Cleidion javanicum is thus the correct name for the species.

            2. Part of continental Asian material is excluded here from C. javanicum and tentatively placed to the closely resembling species C. alongense Bennet & Subh.Chandra (type n.v.). This taxon is characterized by on average larger leaves with coarser teeth, densely hairy staminate rachises, 3-locular ovaries and 3-seeded fruits. It also seems to be mostly geographically apart from C. javanicum, occurring in mountainous areas at NE India, Nepal, Bhutan, China (Yunnan) and northern Burma. With this circumscription C. javanicum has almost exclusively 2-locular ovaries and 1- or 2-seeded fruits (only two specimens seen with perfect 3-seeded fruits).

    Diagnostic characters — The only Malesian species with domatia on lower leaf surface and a predominantly 2-locular ovary.

2. Cleidion luziae Kulju

    Cleidion luziae Kulju in Kulju & Welzen, Blumea 50 (2005) 206, fig. 1a, 3; map 2. — Type: Kajewski 2305 (holo A; iso BM, BRI), Papua New Guinea, Bougainville, Karngu, Buin.

(Small) tree, up to 15 m high, presumably monoecious; flowering branches 1.5–2 mm thick. Bark smooth, grey to brown. Indumentum sericeous, in staminate parts also hirsute and variably oriented, vegetative parts glabrous (to sparsely hairy on stipules outside), generative parts (sub)glabrous to sparsely hairy (ovary densely hairy); granulate surfaces absent. Stipules 1.5–3 by 0.75–1 mm, caducous (to persistent). Leaves inserted evenly along branches or in Terminalia-growth; petiole 0.8–4.6 mm long, sometimes with 2 raised glands at apex; blade (ovate to) elliptic, 8–20 by 2.7–7.3 cm, length/width ratio 2.4–3.4, base rounded to acute, margin subentire to finely serrate, teeth 3–14 per side, apex acuminate, very apex acute, lower surface without domatia; nerves 6–9 per side. Staminate inflorescences: only small fragments seen, basally c. 0.5 mm thick, flowers 1 per node; bracts 0.25–0.5 by 0.2–0.5 mm. Staminate flowers: buds 1.25–1.5 mm diam., brown when dry; sepals c. 1.75 by 1 mm; stamens 59–76, filaments up to c. 1 mm long, anthers 0.3–0.4 by 0.4–0.5 mm, apiculum c. 0.1 mm long. Pistillate inflorescences 3.5–6 cm long, basally 0.5–1 mm thick, 1-flowered, with 2–5 sterile bracts; latter 1–2.5 by 0.5–1.25 mm, persistent. Pistillate flowers: pedicel usually with abscission zone; sepals (3 or) 4 or 5, 1–2 by 0.75–1.5 mm, persistent; locules 3; style 1.5–5 mm long; stigmas erect, 13–24 mm long. Fruits usually 3-seeded, 8–10 by 13–16 mm, ± smooth, brown when dry. Seeds 7–7.5 by 7–8 mm; hilum subcircular, somewhat raised, 1–1.5 by c. 1 mm.

    Distribution — Moluccas (Buru, Seram), New Guinea, Solomon Islands.

    l = C. luziae; « = C. moniliflorum

    Habitat & Ecology — Forests, often on limestone. Altitude up to 1000 m. Flowering and fruiting: mainly September to December.

    Vernacular names — Moluccas: Ai batu, buah hijau muda, daun panjang lancip, perbukitan, tumbuh banyak di batu-batu/jurung (Seram Isl.). Papua New Guinea: Nu-marrie (Bougainville).

    Uses — The fruits are eaten with betel nut to relieve colds (Bougainville).

    Note — This new species, represented by 5 specimens, resembles C. javanicum, but lacks domatia and has single-flowered staminate nodes, persistent pistillate bracts, always 3-locular ovaries and smaller seeds. In addition, the pistillate pedicels of C. luziae usually have an abscission zone. Vietnamese C. bracteosum Gagnep. also has staminate inflorescences with single-flowered nodes, but this species differs from C. luziae in having broader, subcoriaceous leaves and much longer staminate bracts.

3. Cleidion microcarpum Merr.

    Cleidion microcarpum Merr., Philipp. J. Sci. 30 (1926) 404; Kulju & Welzen, Blumea 50 (2005) 208, fig. 4, map 3. — Type: BS (Ramos & Edaño) 44226 (holo UC; iso A, B, K, US), Philippines, Sulu Archipelago, Tawi Tawi.

Shrub to small tree, up to 5(–10) m high, dbh up to 10 cm, mono- or dioecious; flowering branches 1–1.5 mm (staminate) or 1.5–2 mm (pistillate) thick. Bark grey, slightly roughened. Indumentum sericeous, vegetative parts (subglabrous to) sparsely hairy (except leaf blade), generative parts (subglabrous to) sparsely hairy (ovary densely hairy); granulate surfaces present. Stipules 2.5–3.5 by 0.75–1.25 mm, early caducous. Leaves inserted evenly along branches; petiole 0.8–2.5 cm long, apical glands absent; blade elliptic (to some slightly obovate), 6.6–15 by 2.4–6.5 cm, length/width ratio 2–2.8(–3.5), base acute to cuneate to slightly attenuate, margin subentire to crenate (to serrate), teeth 8–16 per side, apex (rounded to) acuminate, very apex (truncate to) rounded, upper surface (sub)glabrous, lower surface (subglabrous to) sparsely hairy only on midrib and nerves, domatia absent; nerves 6–9 per side. Staminate inflorescences up to 7 cm long, basally 0.5–0.75 mm thick, flowers several per node, in fascicles; bracts 0.75–1 by 0.75–1 mm. Staminate flowers: pedicels up to 1 mm long; buds 1–1.5 mm diam., light brown when dry, without a distinctly granulate surface; sepals 1.5–2 by 1–1.5 mm; stamens 50–58, filaments up to 0.75 mm long, anthers 0.3–0.4 by 0.4–0.6 mm, apiculum less than 0.1 mm long. Pistillate inflorescences 4.5–11 cm long, basally 0.5–0.75 mm thick, with 1–5 nodes also in basal half, 1 flower per node; bracts 0.75–1 by 0.75–1 mm, persistent. Pistillate flowers: pedicel 20–35 mm long, with abscission zone in middle to apical part (7–23 mm from base); sepals 3, 1–1.5 by 0.5–1 mm, without distinctly granulate surface, persistent; locules 3; style c. 0.25 mm long; stigmas erect, 3–8 mm long. Fruits 5–6 by c. 8 mm, readily opening, ± smooth, brown when dry. Seeds 3.5–4 by 3.5–4; hilum ± elliptic, 0.5–0.75 by 0.1–0.25 mm.

    Distribution — Philippines (Palawan, Balabac, Ticao, Cebu, Tawi Tawi).

  l = C. microcarpum; «= C. ramosii; n = C. minahassae

    Habitat & Ecology — (Disturbed and secondary) forests, often on limestone. Altitude up to 670 m. Flowering and fruiting: mainly May to October.

    Vernacular names — Philippines: Piligaw-bukid, sarum puti, yambon-yambon (Tagbanua).

    Diagnostic characters — This species may be difficult to distinguish vegetatively from C. ramosii var. ramosii, which also occurs in the Philippines. Although the measurements of these species overlap, C. microcarpum tends to have shorter petioles and smaller leaves with a mainly crenate (as opposed to a mainly serrate) margin.

4. Cleidion minahassae Pax & K.Hoffm.

    Cleidion minahassae Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv (1919) 23; Airy Shaw, Kew Bull. 37 (1982) 12; Kulju & Welzen, Blumea 50 (2005) 210, fig. 5, map 3. — Type: Warburg 15544 (B†), Celebes, Minahassa. Neotype (designated by Kulju & Welzen, 2005): Forsten 11 (holo L, barcode 0158342; iso L, barcodes 0158343, 0158346, 0158349), Celebes, in sylvis Tondano.

Tree; flowering branches 2–4 mm (staminate) or 1.5–2 mm (pistillate) thick. Indumentum sericeous, vegetative parts (subglabrous to) sparsely hairy (except leaf blade), generative parts (subglabrous to) sparsely hairy (ovary densely hairy); granulate surfaces present. Stipules 3–5 by 1.5–2, caducous to somewhat persistent. Leaves inserted evenly along branches; petiole 1.5–6.5 cm long, apical glands absent; blade elliptic to some slightly obovate, 7.5–23 by 3.6–10.8 cm, length/width ratio 2.1–2.9, base acute to cuneate to slightly attenuate, margin serrate, teeth 12–24 per side, apex acuminate, very apex truncate to acute, upper surface (sub)glabrous, lower surface sparsely hairy only on midrib and nerves, domatia absent; nerves 9–13 per side. Staminate inflorescences up to 15 cm long, basally 1–1.5 mm thick, flowers many per node, in glomerules; bracts c. 1 by 1 mm. Staminate flowers: mature flowers not seen; buds dark brown when dry, without a distinctly granulate surface; stamens 78–99. Pistillate inflorescences 10–14 cm long, basally 0.75–1.25 mm thick, with 2–14 nodes also in basal half, 1 or 2 flowers per node; bracts c. 1.25 by 1 mm, persistent. Pistillate flowers: pedicel 7–30 mm long, with abscission zone in middle to apical part (5–20 mm from base); sepals 3 or 4, 1.5–2 by 0.5–1 mm, without distinctly granulate surface, persistent; locules 3; style 0.25–0.5 mm long; stigmas erect, 6–8 mm long. Fruits and seeds not seen.

    Distribution — Indonesia: north-eastern Sulawesi.

  l = C. microcarpum; «= C. ramosii; n = C. minahassae

    Note — The original type (Warburg 15544) in B has been destroyed. The description here is based on four sheets at L (Forsten 11, Tondano). These sheets, presumably representing a single collection, match the original diagnosis in most parts well and are here designated as a neotype.

5. Cleidion moniliflorum Airy Shaw

    Cleidion moniliflorum Airy Shaw, Kew Bull. 34 (1980) 591; Kew Bull. Add. Ser. 8 (1980) 58; Kulju & Welzen, Blumea 50 (2005) 211, fig. 1c, map 2. — Cleidion javanicum Blume var. moniliflorum (Airy Shaw) Chakrab. & M.Gangop., J. Econ. Taxon. Bot. 12 (1988) 493. — Cleidion spiciflorum (Burm.f.) Merr. var. moniliflorum (Airy Shaw) Radcl.-Sm. & Govaerts, Kew Bull. 52 (1997) 478. — Type: NGF (Henty & Frodin) 27334 (holo K; iso L), New Britain, Wariai subdist., Upper Pulie River, near Benim Village.

Tree, monoecious; flowering branches 3–9 mm (staminate) or c. 2 mm (pistillate) thick. Indumentum sericeous, vegetative parts (subglabrous to) sparsely hairy (except leaf blade), generative parts (subglabrous to) sparsely hairy; granulate surfaces present. Stipules 3–5 by c. 1 mm, caducous. Leaves inserted evenly along branches; petiole 0.5–2 cm long, apical glands absent; blade elliptic to some slightly obovate, 9–20 by 3.5–6.5 cm, length/width ratio 2.6–3.5, base acute to cuneate (to slightly attenuate), margin (subentire to) finely serrate, teeth 19–29 per side, apex acuminate, very apex acute, upper surface (sub)glabrous, extra-floral nectaries absent, lower surface (subglabrous to) sparsely hairy only on midrib and nerves, domatia absent; nerves 11–14 per side. Staminate inflorescences ramiflorous, often 2 together, up to 50 cm long, basally c. 1 mm thick, flowers many per node, in glomerules; bracts c. 1 by 1 mm. Staminate flowers: mature flowers not seen; buds dark brown to dark grey when dry, granulate; stamens c. 40. Pistillate inflorescences: only fragment of premature inflorescence seen, with at least 3 nodes. Pistillate flowers, fruits and seeds not seen.

    Distribution — Papua New Guinea: New Britain (only known from type locality).

    l = C. luziae; « = C. moniliflorum

    Habitat & Ecology — Forest, undulating terrain. Altitude 120 m. Flowering: March.

    Note — It is clearly erroneous to treat this species as a variety of C. javanicum (Chakrabarty & Gangopadhyay, 1988), because it differs from C. javanicum in several critical characters (hairs in vegetative parts present, surfaces granulate, staminate flowers in glomerulate clusters, pistillate inflorescences with several flowers).

6. Cleidion papuanum Lauterb.

    Cleidion papuanum Lauterb. in K.Schum. & Lauterb., Nachtr. Fl. Schutzgeb. Südsee (1905) 296; Pax &  K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 293; Whitmore, Guide Forests Brit. Solomon Isl. (1966) 173; Airy Shaw, Kew Bull. 23 (1969) 85; Kew Bull. Add. Ser. 8 (1980) 58; Kulju & Welzen, Blumea 50 (2005) 212, fig. 1d, fig. 6a–c, map 4. — Type: Schlechter 14122 (holo WRSL; iso BM, BO, P), Kaiser-Wilhelmsland [= Papua New Guinea], Weg vom Ramu zur Küste.

    Trigonostemon oliganthum K.Schum. in K.Schum. & Lauterb., Nachtr. Fl. Schutzgeb. Südsee (1905) 298. — Type: Nyman 554 (UPS), Kaiser-Wilhelmsland [= Papua New Guinea], Sattelberg bei 800 m.

    Cleidion salomonis Airy Shaw, Kew Bull. 23 (1969) 87. — Type: BSIP (Whitmore) 4233 (holo K; iso L), Solomon Islands, San Cristobal, Wairaha River, 5 miles from north coast. Syn.nov.

Shrub to small tree, usually few-branched, up to 5(–12) m high, dbh up to 8 cm; (mono- or) dioecious; flowering branches 1.5–10 mm thick. Bark yellow to brown to grey, smooth (to fissured). Indumentum sericeous to hirsute (erect or variably oriented), vegetative parts sparsely (to densely) hairy (except leaf blade), generative parts (subglabrous to) sparsely (to densely) hairy; granulate surfaces present. Stipules 4–45 by 0.5–5 mm, caducous to fairly persistent. Leaves well separated or ± tightly grouped at the branch ends; petiole 0.4–2.5 cm long, apical glands absent; blade obovate, 9.8–73 by 2.5–17 mm, length/width ratio 2.8–7.5, base gradually narrowing, cuneate (to attenuate), margin (crenate to) serrate, basally often subentire, teeth 26–80 per side, apex (obtuse to) acuminate, very apex (rounded to) acute to acuminate, upper surface glabrous to sparsely hairy on venation, lower surface (subglabrous to) sparsely hairy, hairs mainly on venation, domatia absent; nerves 11–34 per side. Staminate inflorescences up to 120 cm long, basally 0.75–4 mm thick, flowers many per node, in glomerules or often slightly elongated, branching clusters; bracts 0.75–2 by 0.5–1.5 mm. Staminate flowers: pedicels up to 2.5 mm long; buds 1–1.5 mm diam., brown to dark grey when dry, without distinctly granulate surface; sepals c. 1.5 by 1 mm; stamens 45–67, filaments up to 1 mm long, anthers 0.3–0.5 by 0.4–0.6 mm, apiculum less than 0.1 mm long. Pistillate inflorescences 10–45 cm long, basally 0.75–2 mm thick, with 2–6(–16) nodes in upper half, 1(–3) flowers per node; bracts 1–4 by 0.5–1.5 mm, persistent. Pistillate flowers: pedicel 10–25 mm long, with abscission zone at the basal to apical part (1–22 mm from base); sepals 3, 1–4 by 0.5–1.5 mm, persistent; locules 3 (or 4); style 0–1.5 mm long; stigmas erect to slightly ascending, 7–20 mm long. Fruits c. 6 by 10 mm, readily opening, surface ± smooth, green to cream to red when fresh, brown when dry. Seeds c. 6 by 5 mm; hilum elliptic, c. 1 by 0.75 mm.

    Distribution — New Guinea, Solomon Islands.

    Habitat & Ecology — Primary (to secondary) rain forests, often on ridges and hillsides, well to poorly drained soils. Altitude up to 900 m. Flowering and fruiting all year round.

    Vernacular names — Papua New Guinea: Fokko (Manikiong). Solomon Islands: Sa’a, saola (Kwara’ae).

    Note — Cleidion salomonis, a species with large leaves, is synonymised with C. papuanum. The type of the former represents one end of a continuum in a vast variation in leaf and stipule size. The specimens with small or large leaves are also geographically intermixed.

    Diagnostic characters — Only Malesian species with always clearly obovate leaves and gradually narrowing leaf base.

7. Cleidion ramosii (Merr.) Merr.

    Cleidion ramosii (Merr.) Merr., Philipp. J. Sci. 20 (1922) 400; Enum. Philipp. Fl. Pl. 2 (1923) 439; Airy Shaw, Alph. Enum. Euphorb. Philipp. Isl. (1983) 13; Kulju & Welzen, Blumea 50 (2005) 214, fig. 6d–g, map 3. — Mallotus ramosii Merr., Philipp. J. Sci., Bot. 7 (1912) 401; Pax &  K.Hoffm. in Engl., Pflanzenr. IV.147.vii (1914) 195. — Lectotype (designated by Kulju & Welzen, 2005): BS (Ramos) 14602 (holo US; iso A, K), Philippines, Camiguin de Mindanao, near the summit of Mt Catarman.

    Cleidion lanceolatum Merr., Philipp. J. Sci., Bot. 9 (1914) 474; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv (1919) 23; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 439; Airy Shaw, Alph. Enum. Euphorb. Philipp. Isl. (1983) 13. — Lectotype (designated by Kulju & Welzen, 2005): Ramos 1648 (holo G; iso A, BM, L, NSW, NY), Philippines, Samar. Syn. nov.

    Mallotus samarensis Merr., Philipp. J. Sci., Bot. 9 (1914) 488. — Type: BS (Ramos) 17480 (iso K, US), Philippines, Samar, Yabong.

Shrub to small tree, up to 8 m high, dbh up to 13 cm, monoecious; flowering branches 1–3.5 mm thick. Indumentum sericeous, vegetative parts (subglabrous to) sparsely hairy (except leaf blade), generative parts (subglabrous to) sparsely hairy (ovary densely hairy); granulate surfaces present. Stipules 2.5–5 by 1–2 mm, caducous to somewhat persistent. Leaves inserted evenly along branches; petiole 0.8–8 cm long, apical glands absent; blade (ovate to) elliptic (to some slightly obovate), 7.8–23 by 1.4–9.3 cm, length/width ratio 1.9–8.8, base (rounded to) acute to cuneate to attenuate, margin subentire (to crenate) to serrate, teeth 9–23 per side, apex acuminate, very apex truncate to acute, upper surface (sub)glabrous, lower surface (subglabrous to) sparsely hairy only on midrib and nerves, domatia absent; nerves (7–)8–14 per side. Staminate inflorescences up to 24 cm long, basally 0.5–1.25 mm thick, flowers many per node, in glomerules; bracts 1–2.5 by 1–2 mm. Staminate flowers: pedicels up to 3 mm long; buds 1.25–2 mm diam., dark brown to dark grey when dry, granulate; sepals c. 2 by 1.5 mm; stamens 60–108, filaments up to c. 1 mm long, anthers 0.2–0.4 by 0.3–0.6 mm, apiculum up to 0.1 mm long. Pistillate inflorescences 6–18(–27) cm long, basally 0.5–0.75 mm thick, with 1–5 nodes in upper half, 1 (or 2) flowers per node; bracts 1.25–2.5 by 1–1.5 mm, persistent. Pistillate flowers: pedicel 4–23 mm long, with abscission zone in basal part (1–9 mm from base); sepals 3–5, 1.5–3 by 0.5–1.5 mm, usually distinctly granulate, persistent; locules 3; style absent to up to 0.5 mm long; stigmas erect to ascending, 4.5–13 mm long. Fruits 4–5 by c. 8 mm, readily opening, ± smooth, brown when dry. Seeds 3–4.5 by 3.5–4.5 mm; hilum ± elliptic, 0.5–1 by 0.1–0.5 mm.

    Distribution — Philippines.

  l = C. microcarpum; «= C. ramosii; n = C. minahassae

    Habitat & Ecology — Primary and secondary forests; often along rivers and on limestone. Altitude up to 470 m. Flowering and fruiting all year round.

    Note — Cleidion lanceolatum is treated here as a rheophytic variety of C. ramosii, leaf length/width ratio being the only unambiguous character to separate the taxa. Variety lanceolatum also tends to have somewhat longer stipules, bracts, pistillate pedicels, sepals and stigmas, but these measurements overlap with those of var. ramosii.

    Diagnostic characters — See under C. microcarpum.

Key to the varieties

a. var. ramosii

 

    Cleidion ramosii (Merr.) Merr. var. ramosii: Kulju & Welzen, Blumea 50 (2005) 215.

(for nomenclature see C. ramosii and M. samarensis under the species)

 

Leaves: petiole 1–8 cm long; blade (ovate to) elliptic (to slightly obovate), 7.8–23 by 3.2–9.3 cm, length/width ratio 1.9–3; nerves (7–)8–11 per side.

    Distribution — Philippines (Mindoro, Tablas, Samar, Siargao, Mindanao).

 

b. var. lanceolatum (Merr.) Kulju

 

    Cleidion ramosii (Merr.) Merr. var. lanceolatum Kulju in Kulju & Welzen, Blumea 50 (2005) 215. — Cleidion lanceolatum Merr. Philipp. J. Sci., Bot. 9 (1914) 474; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xiv (1919) 23; Merr., Enum. Philipp. Fl. Pl. 2 (1923) 439; Airy Shaw, Alph. Enum. Euphorb. Philipp. Isl. (1983) 13. — Lectotype (designated by Kulju & Welzen, 2005): Ramos 1648 (holo G; iso A, BM, L, NSW, NY), Philippines, Samar.

 

Leaves: petiole 0.8–3 cm long; blade narrowly to linear elliptic, 9.5–19.8 by 1.4–3.6 cm, length/width ratio 3.4–8.8; nerves 9–14 per side.

    Distribution — Philippines (Samar, Bohol).

    Habitat & Ecology — Often along rivers.

Doubtful species

    Cleidion megistophyllum (Quisumb. & Merr.) Airy Shaw, Kew Bull. 25 (1971) 528; Alph. Enum. Euphorb. Philipp. Isl. (1983) 13; Kulju & Welzen, Blumea 50 (2005) 215. — Actephila megistophylla Quisumb. & Merr., Philipp. J. Sci. 37 (1928) 158. — Lectotype (designated by Kulju & Welzen, 2005): BS (Ramos & Edaño) 45466 (holo A; iso K, NY, US), Philippines, Luzon, Tayabas Prov., Casiguran.

Indumentum on pistillate parts sericeous, (subglabrous to) sparsely hairy, leaves glabrous; granulate surfaces present. Stipules not seen. Leaves: petiole 2–3.5 cm long, thick; blade obovate, 115–155 by 21–26 cm, length/width ratio 4.4–7.4, base cuneate, margin finely serrate, teeth c. 40 per side, apex acute; upper and lower surfaces without extra-floral nectaries, domatia absent; nerves 40–45 per side. Staminate inflorescences and flowers not seen. Pistillate flowers not seen. Infructescences c. 12 cm long, basally 3–4 mm thick, with 3–8 fruits, 1 flower per node; bracts c. 5 mm wide, caducous to persistent; pedicels 4–9 cm long, with 3–6 bracteoles at basal part, abscission zone absent; sepals 5, ovate to obovate, 3–6 by 2.5–4 mm, apex rounded to obtuse; within the sepals remnants of a darker coloured seemingly disc-like structure surrounding the column and splitting into 1–2 mm long segments. Fruits: not seen completely, presumably 3-seeded, readily opening, ± smooth, dark brown to dark grey when dry, speckled. Seeds 17–19 by 17–19 mm; hilum 1–2 by 0.5–1 mm.

    Distribution — Philippines (Luzon).

    Notes — 1. This species is known only from an incomplete type collection (including only leaves and remnants of infructescences and fruits). According to the original description (Quisumbing & Merrill, 1928), it is an up to 4 m high shrub occurring in damp forests along streams at low altitudes.

2. The absence of both staminate and pistillate flowers makes it impossible to indisputably confirm the placement of this species in genus Cleidion. The absence of extra-floral nectaries on both leaf surfaces, the presence of disc-like remnants inside the pistillate sepals, and the unusually long (relative to infructescence size) pistillate pedicels are deviating characters from all other (Malesian) Cleidion species.

Excluded species

    Acalypha spiciflora Burm.f., Flora Indica (1768) 203. — Cleidion spiciflorum (Burm.f.) Merr., Interpr. Herb. Amboin. (1917) 322, in obs., pro typo. — Type: Herb. Burman (holo G, n.v., photo seen) = Claoxylon sp. (indicated by Merrill’s note accompanying the type).

    Cleidion bishnui Chakrab. & M.Gangop., J. Econ. Taxon. Bot. 12 (1988) 473 = Cleidiocarpon laurinum Airy Shaw