Malesian Euphorbiaceae Descriptions |
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Widuri, R. & P.C. van Welzen. 1998. A revision of the genus Cephalomappa (Euphorbiaceae) in Malesia. Reinwardtia 11: 153184.
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Cephalomappa Baill., Adansonia 2 (1874) 131; Pax in Engl., Pflanzenr. IV.147.ii (1910) 16; Pax & K.Hoffm. in Engl. & Harms, Nat. Pflanzenfam. 2 ed., 19c (1931) 123; Airy Shaw, Kew Bull. 14 (1960) 380; Kew Bull. 16 (1963) 353; Whitmore, Tree Fl. Malaya 2 (1973) 76; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 66; Airy Shaw, Kew Bull. 36 (1981) 274; G.L.Webster, Ann. Missouri Bot. Garden 81 (1994) 79; Widuri & Welzen, Reinwardtia 11 (1998) 161; Radcl.-Sm., Gen. Euphorbiacearum (2001) 186; G.L.Webster in Kubitzki, Fam. Gen. Vasc. Pl. 11 (2014) 137. Type: Cephalomappa beccariana Baill.
Trees, monoecious. Indumentum: next to simple hairs usually with lepidote and stellate hairs. Branches smooth to rough. Stipules small, linear, very early caducous. Leaves alternate, simple; petiole terete, basally and apically pulvinate, usually hairy; blade ovate to obovate, symmetric, coriaceous; base rounded-obtuse or acute; margin entire to crenate to dentate, flat or recurved, with glands on the lower surface, apex acuminate to cuspidate, very apex obtuse to acute, upper surface glabrous, sometimes with stellate hairs on the basal part of the midrib, lower surface hairy or glabrous; venation pinnate, raised on both sides, especially below, nerves 411 pairs, submarginally looped, united, veins scalariform; glands 14 basally beneath. Inflorescences axillary, thyrsoid, paniculate, with the staminate flowers in glomerules and the pistillate flowers solitary; bracts ovate, with stellate hairs, apex (obtusely) acute; bracteoles ovate or obovate; peduncles and pedicels hairy or glabrous. Flowers actinomorphic, sessile or pedicellate. Staminate flowers: calyx 3-merous, campanulate, valvate, lobes rounded or acute, thinly to densely papillate; petals and disc absent; staments 3-5; filaments basally united into a glabrous to hairy androphore; anthers subapically dorsifixed, opening latrorsly, lengthwise, smooth; pistillode small. Pistillate flowers: calyx either connate and 5-lobed or sepals free, 8, angle between lobes obtuse or acute, apex entire or bifid, acute or obtuse, with a gland; ovary echinate to densely echinate, hairy, 3- or 4-locular; ovules 1 per locule; style short to long, terete, glabrous or with stellate hairs; stigmas 3 or 4, apically entire or bifid, papillate above, smooth below. Fruit a rhegma, ellipsoid in lateral view, 3-lobed, all lobes developed, echinate to densely echinate, with stellate hairs or glabrous; wall thick, woody, smooth and glabrous inside; peduncle hairy or galbrous. Seed subglobular, smooth, glabrous, apex 2-lobed; hilum ovate. Leaf anatomy: abaxial surface of leaf hairy or glabrous; epidermal anticlinal walls sinuous to deeply sinuous anticlinal or almost straight to sligtly wavy; stomata paracytic, elliptic.
Distribution Five species are recognized, all in Malesia: Malay Peninsula, Sumatra, and Borneo.
1a. |
Branches rough. Lower leaf suface hairy. |
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1b. |
Branches smooth. Lower leaf surface glabrous. |
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Androphore subglabrous. Stigma apically entire. Fruit echinate. |
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2b. |
Androphore glabrous. Stigma apically bifid. Fruit densely echinate. |
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Leaves ovate or elliptic. Stamens 3. |
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3b. |
Leaves elliptic. Stamens 4 (or 5). |
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Leaves obovate, margin serrate. Staminate bracteoles with a fimbriate apex. Pistillate sepals connate into a calyx; stigmas 3. |
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4b. |
Leaves elliptic, margin crenate. Staminate bracteoles with an acute apex. Pistillate sepals free; stigmas 4. |
Cephalomappa beccariana Baill., Adansonia 2 (1874) 131; Pax in Engl., Pflanzenr. IV.147.ii (1910) 17; Airy Shaw, Kew Bull. 14 (1960) 380; Kew Bull. Add. Ser. 4 (1975) 66; Widuri & Welzen, Reinwardtia 11 (1998) 163, Fig. 48, Map 1. Type : Beccari PB 425 (holo FI, n.v.; iso in K), Sarawak.
Tree. Branches rough, flowering branches c. 3.5 mm thick. Leaves: petiole 1.13.5 cm long, covered by dense long and short stellate hairs; blade ovate to elliptic, 420.2 by 29.3 cm, base obtuse to rounded, with a single gland at the lower surface close to the midrib, margin entire to subentire to shallowly denticulate, flat, apex acuminate to cuspidate, very apex acute, upper surface glabrous except for basal part of midrib, lower surface covered with dense short lepidote to sparse (tuffed) short to dense long and short stellate hairs, nerves 57 pairs. Inflorescences: peduncle 23.5 cm long, densely hairy; peduncle to glomerules 0.12.7 cm long, densely hairy, glomerules 25 by c. 2.6 mm; staminate bracts 0.81.2 by 0.20.4 mm; staminate bracteoles 1.21.4 by 0.20.4 mm, margin fimbriate; pistillate bracts 11.4 by 0.20.4 mm, apex obtuse, with stellate hairs. Staminate flowers 2428 together, 1.63.4 by 0.11.2 mm; calyx tube 1.22.4 by 0.81.6 mm; lobes 0.50.8 b 0.30.5 mm, subglabrous, densely papillate; stamens 3, androphore 0.30.5 by 0.150.3 mm, subglabrous to with few stellate hairs, filaments 1.62.4 mm long, anther 0.60.8 by 0.30.4 mm; pistillode 12.2 mm long. Pistillate flowers: pedicels 0.23.1 mm long, with short lepidote hairs or long and short stellate hairs; calyx 5-lobed, with dense long and short stellate hairs; tube 0.31.7 by 0.91.8 mm; lobes 0.81.3 by 0.41.2 mm; ovary 0.92.3 by 0.62.0 mm, style 0.61.6 mm long; stigmas 3, 0.71.5 mm long, apically entire, surface papillate. Fruits 1.21.9 by 1.32.6 cm, echinate, with long and short stellate hairs, peduncle 24.5 cm by 1.42.5 mm; wall 2.73 mm thick. Seeds 0.71.3 by 0.71.3 cm. Leaf anatomy: leaf abaxially with dense short (7590 ΅m long) lepidote hairs, or sparse to dense long (125163 ΅m) and short (6394 ΅m) stellate hairs; epidermis cells with slightly wavy to (deeply) sinuous anticlinal walls, 3644 by 1829 ΅m wide. Stomata 2124 by 1618 ΅m.
Distribution Malesia : Borneo (Sarawak).
1a. |
Lower leaf surface with stellate hairs; epidermis cells with sinuous anticlinal walls. |
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1b. |
Lower leaf surface with lepidote hairs; epidermis cells with slightly wavy anticlinal walls. |
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2a. |
Blade coriaceous, margin entire. |
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2b. |
Blade papery, margin subentire to shallowly denticulate. |
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3a. |
Leaves ovate, lower surface with dense stellate hairs, epidermis cells with deeply sinuous anticlinal walls. |
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3b. |
Leaves elliptic, lower surface with sparse stellate hairs, epidermis cells with sinuous anticlinal walls. |
var. beccariana
Cephalomappa beccariana Baill. var. beccariana: Airy Shaw, Kew Bull. 14 (1960) 380; Kew Bull. Add. Ser. 4 (1975) 66; Widuri & Welzen, Reinwardtia 11 (1998) 165, Fig. 5. Type : as the species.
Leaves: petioles 1.23.5 cm long; blade ovate, 612 by 45.8 cm, coriaceous, base rounded, margin entire, lower surface covered with dense long and short stellate hairs. Peduncle to staminate glomerules 1.11.7 cm long, densely hairy; glomerules 4.55 by 2.63.1 mm. Staminate flowers 2.93.4 by 0.81.2 mm; calyx: tube 2.22.4 by 0.81.2 mm, lobes 0.70.8 by 0.30.5 mm, subglabrous; stamens: androphore 0.30.5 by 0.20.3 mm, filaments 1.82.4 mm long; pistillode 1.62 mm long. Pistillate flowers: pedicels 13.1 cm long, covered with long and short stellate hairs; calyx: tube 0.30.5 by 1.41.8 mm, lobes 1.11.3 by 0.61.2 mm; ovary 1.92.3 by 1.62 mm, style 1.21.6 mm long, stigmas 1.31.5 mm long. Fruits 1.51.9 by 2.42.6 cm. Seeds 1.21.3 by 1.11.3 cm. Leaf anatomy: abaxial epidermis with dense long (125163 ΅m) and short (6394 ΅m) stellate hairs; epidermis cells with deeply sinuous anticlinal walls, 4143 by 2426 ΅m; stomata c. 24 by 18 ΅m.
Distribution Malesia: Borneo (Sarawak).
Habitat & Ecology Primary lowland forest, on hill slopes with yellow sandy soil. Alt: sea level up to 275 m.
Vernacular names Sarawak: Mahu hutan (Malay).
Uses The fruit is locally eaten.
Note Baillon (1874) described the apex of the stigmas as bifid. We found those to be entire.
var. havilandii Airy Shaw
Cephalomappa beccariana Baill. var havilandii Airy Shaw, Kew Bull. 14 (1960) 380; Kew Bull. 16 (1963) 353; Kew Bull. Add. Ser. 4 (1975) 66; Widuri & Welzen, Reinwardtia 11(1998) 167, Fig. 6. Type: Haviland 2184 (K, holo, n.v.; iso BM, L, BO), Sarawak, near Kuching.
Leaves: petioles 1.42.3 cm long; blade elliptic, 48 by 24.3 cm, coriaceous, base obtuse; margin entire, lower surface with dense short lepidote hairs. Peduncle to staminate glomerules 25 mm long with short lepidote hairs; glomerules 24 by 46 mm. Staminate flowers 1.61.8 by 0.81 mm; calyx tube 1.21 by 1.216 mm, lobes 0.50.7 by 0.30.5 mm, subglabrous; stamens: androphore 0.30.5 by 0.20.3 mm, subglabrous, filaments 1.62 mm long; pistillode 1.82.2 mm long. Pistillate flowers: sessile or pedicelled, latter with short lepidote hairs; calyx: tube 0.30.5 by 1.31.5 mm, lobes 0.81.2 by 0.40.6 mm; ovary 1.92.1 by 1.61.8 mm, style 1.21.3 mm long, stigmas 0.91.3 mm long. Fruits 1.51.7 by 1.31.5 cm. Seeds 0.81 by 0.7-0.9 cm. Leaf anatomy: abaxial epidermis with dense short (7590 ΅m long) lepidote hairs; epidermis cells with slightly wavy anticlinal walls, 3642 by 1822 ΅m; stomata c. 22 by 16 ΅m.
Distribution Malesia: Borneo (Sarawak).
var. hosei Airy Shaw
Cephalomappa beccariana Baill. var. hosei Airy Shaw, Kew Bull. 14 (1960) 380; Kew Bull. Add. Ser. 4 (1975) 66; Widuri & Welzen, Reinwardtia 11 (1998) 168, Fig. 7. Type: Hose 303 (K, holo, n.v.; iso in L), Sarawak, Baram.
Leaves: petioles 1.13.2 cm long; blade elliptic, 618 by 47.4 cm, coriaceous, base rounded, margin entire, lower surface with sparse short stellate hairs. Peduncle to staminate glomerules (0)13.1 mm long, covered with dense short stellate hairs; glomerules c. 4 by 3.75 mm. Staminate flowers c. 1.9 by 0.9 mm; calyx tube c. 1.2 by 0.9 mm, lobes c. 0.7 by 0.4 mm; stamens: androphore c. 0.3 by 0.15 mm subglabrous, filaments c. 1.8 mm long; pistillode 12 mm long; Pistillate flowers sessile or with 13.1 mm long pedicels, the latter covered with dense short stellate hairs; calyx tube 1.51.7 by 1.41.6 mm, lobes 0.91.2 by 0.50.7 mm; ovary 1.82 by 1.71.9 mm, style 1.21.4 mm long, stigma 0.91.1 mm long. Fruits 1.41.6 by 1.92.3 cm. Seeds 0.70.9 by 0.80.9 cm. Leaf anatomy: abaxial epidermis with sparse short (6094 ΅m long) stellate hairs. Epidermis cells with sinuous anticlinal walls, 4244 by 2529 ΅m; stomata c. 23 by 16 ΅m.
Distribution Malesia: Borneo (Sarawak).
var. tenuifolia Airy Shaw
Cephalomappa beccariana Baill. var. tenuifolia Airy Shaw, Kew Bull. Add. Ser. 4(1975) 66; Widuri & Welzen, Reinwardtia 11 (1998) 169, Fig. 8 Type: S 18041 (K, holo; iso in BM, BO), Sarawak, First Division, Lundu District, Gunong Gading.
Leaves: petioles 1.23.5 cm long; lamina elliptic, 820.2 by 59.3 cm, papery, base rounded, margin subentire to shallowly denticulate, lower surface covered with sparse stellate hairs in tufts. Penduncle to staminate glomerules 22.7 cm long, covered with dense short stellate hairs; glomerules c. 3 by 4 mm. Staminate flowers 1.82.2 by 11.1 mm; calyx tube 1.21.4 by 0.91.1 mm, lobes 0.60.8 by 0.30.5 mm; stamens: androphore 0.30.5 by 0.20.3 mm, subglabrous, filaments 1.82.2 mm long; pistillode 1.31.5 mm long. Pistillate flowers: pedicels 0.20.3 mm long, covered with dense stellate hairs; calyx tube 0.30.5 by 0.91.1 mm, lobes 0.81 by 0.70.9 mm; ovary 0.91.2 by 0.60.8 mm, style 0.60.8 mm long, stigma 0.70.9 mm long. Fruits 1.21.4 by 2.42.6 cm. Seeds 0.91.1 by 11.2 cm. Leaf anatomy: abaxial epidermis covered with sparse stellate hairs, 69125 ΅m long; epidermis cells with sinuous: anticlinal walls, 4143 by 2124 ΅m; Somata c. 21 by 16 ΅m.
Distribution Malesia: Borneo (Sarawak).
Cephalomappa lepidotula Airy Shaw, Kew Bull. 14 (1960) 379; Whitmore, Tree Fl. Malaya 2 (1973) 76; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 67; Kew Bull. 36 (1981) 275; Widuri & Welzen, Reinwardtia 11 (1998) 170, Fig. 9, 10, Map 2. Type: Beccari PS 975 (K, holo, n.v.; iso in BM, L), Sumatra, Padang Prov., Sungai Bulu.
Tree. Branches rough, flowering branches c. 3.5 mm thick. Leaves petioles 3.25.2 cm long, covered with dense lepidote hairs; blade elliptic, 6.520.2 by 3.45.2 cm, base obtuse, with 2 glands at the lower surface near the midrib, margin dentate, flat, apex acuminate, very apex acute, upper surface glabrous, lower surface with closely set lepidote hairs,nerves 59 pairs. Inflorescences: peduncle 23.5 cm long, densely hairy; peduncles to glomerules 0.71 cm long, densely hairy; glomerules 3.84.2 by 3.63.8 mm; staminate bracts c. 1.1 by 0.9 mm; staminate bracteoles 0.81 by 0.20.3 mm, margin fimbriate; pistillate bracts ovate, 0.91.1 by 0.82.2 mm; apex acute, with lepidote hairs. Staminate flowers 2735 together, 1.61.8 by 0.91.1 mm; calyx tube 1.11.3 by 0.91.1 mm, lobes 0.40.6 by 0.30.5 mm, densely papillate; stamens 4, androphore 0.30.5 by 0.20.3 mm, glabrous, filaments 1.11.3 mm long; anthers 0.60.8 by 0.40.6 mm; pistillode 0.91.1 mm long. Pistillate flowers: pedicels 1.41.6 mm long, covered with dense lepidote hairs; calyx 5-lobed, with lepidote hairs, tube 0.30.5 by 1.41.6 mm; lobes 11.2 by 0.70.9 mm; ovary 0.91.1 by 1.11.4 mm; style 1.61.8 mm long; stigmas 3, 1.31.5 mm long, apically bifid, surface papillate. Fruits c. 3 by 4.2 cm, densely echinate with 2.53.7 mm long spines, lepidote hairs present; peduncle 12.9 by 1.92.9 mm; wall c. 2.4 mm thick, woody, inside smooth and glabrous. Seeds 11.2 by 0.91.1 cm, smooth, glabrous, apex 2-lobed. Leaf anatomy: leaf abaxially covered with lepidote hairs 87113 ΅m long; epidermis cells with wavy anticlinal walls, 3436 by 2022 ΅m; stomata c. 24 by 17 ΅m.
Distribution Malesia: Malay Peninsula, Sumatra, Borneo.
Habitat & Ecology Swampy forest and on hill slopes. Alt: c. 195 m.
Cephalomappa mallolicarpa J.J.Sm., Bull. Jard. Bot. Buitenz. Ser. 3, 6 (1924) 95; Merr., P1. Elmer. (1929) 160; Airy Shaw, Kew Bull. 14 (1960) 380; Whitmore, Tree Fl. Malaya 2 (1973) 76, Fig. 4; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 67; Kew Bull. 36 (1981) 275; Widuri & Welzen, Reinwardtia 11 (1998) 173, Fig. 11, 12; Map 3. Lectotype (Widuri & Welzen, 1998): Bogor Botanical Garden VIII.F.50 (BO, hololocto), Culta Bogor Botanical Gardens.
Tree. Branches smooth, glabrous, flowering branches c. 4.3 mm thick. Leaves: petiole 0.61.2 cm long, glabrous; blade obovate, 1028.2 by 2.56.2 cm, base acute with 2 glands at the lower surface near the midrib. margin serrate, flat, apex cuspidate, very apex obtuse, both surfaces smooth, glabrous, nerves 811 pairs. Inflorescences: peduncle 23 cm long, glabrous; peduncle to glomerules 0.51.8 cm long, glabrous; glomerules 0.50.7 by 0.40.6 cm; staminate bracts 0.91.1 by 0.20.4 mm; staminate bracteoles 1.41.6 by 0.50.7 mm, margin fimbriate; pistillate bracts 0.9-1:1 by 0.3-0.5 mm, apex obtuse, subglabrous. Staminate flowers 2936 together, 1.82.2 by 0.91.1 mm; calyx tube 1.41.7 by 0.91.1 mm; lobes 0.40.5 by 0.40.6 mm, glabrous, densely papillate; stamens 3 (or 4), androphore 0.30.4 by c. 0.2 mm, glabrous, filaments 1.11.3 mm long; pistillode 0.81 mm iong. Pistillate flowers: pedicels 02 cm long, glabrous; calyx 5-lobed, subglabrous, tube 0.30.5 by 1.51.7 mm, lobes 11.2 by 0.50.7 mm; ovary 1.51.7 by 2.52.7 mm, style c. 1.5 mm long, stigmas 3, 1.82 mm, apically usually bifid, upper surface papillate. Fruits 1.11.3 by 2.22.4 cm, echinate; peduncle 30.5 cm by 22.4 mm; wall 2.83 mm thick, woody, inside smooth and glabrous. Seeds 1.11.3 by 0.91.1 cm. Leaf anatomy: leaf abaxially glabrous; epidermis cells with sinuous anticlinal walls, 4446 by 2630 ΅m; stomata c. 26 by 16 ΅m.
Distribution Malesia: Malay Peninsula, Sumatra, Borneo (not known from Brunei and Sarawak yet).
Habitat & Ecology Primary forest, on hillside with a sandy-loam soil with lime. Alt: 80600 m.
Uses The wood is of low quality, its durability class is V. Nevertheless, local people use it for the construction of their house.
Note The original syntypes, cultivated in Bogor Botanical Garden, were said to be from Mt. Salak in Java, but, this species was never found on this mountain. Up to now the species is not known from Java, therefore, Smith's (1924) suspicion of the Mt. Salak origin of the trees seems justified.
Cephalomappa paludicola Airy Shaw, Kew Bull. 14 (1960) 380; Kew Bull. 16 (1963) 353; Whitmore, Tree Fl. Malaya 2 (1973) 76; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 67; Widuri & Welzen, Reinwardtia 11 (1998) 176, Fig. 13, 14, Map 4. Type: S 9713 (K, holo, n.v.; iso in L), Sarawak, Binatang Distr., Surong trib., Daro Forest Reserve.
Tree. Branches smooth, flowering branches c. 3.2 mm thick. Leaves: petiole 0.50.7 cm
long, glabrous; blade elliptic, 6.51:3 by 25 cm, base acute, with 4 glands
at the lower surface near the midrib, margin
Distribution Malesia: Borneo (Sarawak).
Habitat & Ecology Mainly found in freshwater peat swamp forest, also in primary forest. Alt.: sea level up to 300 m.
Vernacular names Sarawak: Arau, ahrau, pela.
Uses The wood is used as firewood.
Note Distinguishable from the other species by its crenate leaf margin, staminate flowers with ovate bracteoles, and the 8 free pistillate sepals with their obtuse apex.
Cephalomappa penangensis Ridl., Fl. Malay Pen. 3 (1924) 279; Airy Shaw, Kew Bull. Misc. Inf. (1923) 368; Kew Bull. 14 (1960) 379; Kew Bull. 16 (1963) 353; Whitmore, Tree Fl. Malaya 2 (1972) 76; Widuri & Welzen, Reinwardtia 11 (1998) 179, Fig. 15, 16, Map 5. Type: Curtis 3584 (SING, iso), Malay Peninsula.
Tree. Branches rough, strong, flowering branches c. 4 mm thick. Leaves: petiole 1.83.2 cm long, covered with densely stellate hairs; blade elliptic, 14.523.5 by 7.410.4 cm, base obtuse, with 2 glands at the lower surface close to the midrib, margin sinuate, recurved, apex acuminate to cuspidate, very apex acute; upper surface glabrous, glossy; lower surface covered with sparse, long stellate hairs, nerves 79 pairs. Inflorescences: peduncle 22.5 cm long, hairy; peduncle to glomerules 0.41.2 mm long. densely hairy; glomerules 68 by 46 mm; staminate bracts 1.21.4 by 0.60.9 mm; staminate bracteoles 1.51.7 by 0.30.4 mm, margin fimbriate; pistillate bracts 1.21.3 by 0.60.8 mm, apex acute, with stellate hairs. Staminate flowers 2734 together, 2.22.3 by 1.72.3 mm; calyx tube 1.41.6 by 1.72.3 mm, lobes 0.70.8 by 0.61.2 mm, densely papillate; stamens 4 (or 5), androphore 0.30.5 by 0.20.3 mm, subglabrous, filaments 1.92.1 mm long, anthers 0.70.9 by 0.30.5 mm; pistillode 0.20.4 mm long. Pistillate flowers: pedicels c. 0.8 cm long, with stellate hairs; calyx 5-lobed, with dense stellate hairs, tube 1.41.6 by 1.81 mm; lobes 0.30.4 by 0.30.5 mm; ovary c. 2.3 by 2.5 mm, style 1.61.8 mm long, stigmas 3, 11.2 mm long, apically entire, upper surface papillate. Fruits 1.41.8 by 2.22.4 cm, echinate, suglabrous; peduncle 13 cm by 1.32 mm; wall 2.73.1 mm thick. Seeds 11.2 by 1.11.3 cm. Leaf anatomy: leaf abaxially covered with long stellate hairs, 125225 ΅m long; epidermis cells with deeply sinuous anticlinal walls, 4244 by 2628 ΅m. Stomata c. 23 by 17 ΅m.
Distribution Malesia: Malay Peninsula.
Habitat & Ecology Lowland forest. Alt.: c. 90 m.
Note C. penangensis resembles C. beccariana var. tenuifolia, but the latter has thin leaves with a flat denticulate margin. C. penangensis, unlike the other species. has much stouter petioles and stiff recurved leaves with a sinuate margin.