|
Malesian Euphorbiaceae Descriptions |
|
Welzen, P.C. van, Q.D. Nguyen & R.C.K. Chung. 1997. A revision of the introduced species of Manihot (Euphorbiaceae) in Malesia. Rheedea 7: 77–85.
Goto on this page:
Manihot Miller, Gard. Dict. ed. 4, 2 (1754); Adans., Fam. Pl. 2 (1763) 356; Crantz, Inst. Rei Herb. 1 (1766) 166; Pohl, Pl. Bras. Icon. Descr. 1 (1827) 17; Müll.Arg. in DC., Prodr. 15, 2 (1866) 1057; in Mart., Fl. Bras. 11, 2 (1874) 438; Pax in Engl., Pflanzenr. IV.147.ii (1910) 21; Standl., Contr. U.S. Natl. Herb.23 (1923) 642; Pax & K.Hoffm. in Engl., Pflanzenr. IV.147.xvi (1924) 194; Croizat, J. Arnold Arbor. 23 (1942) 216; Backer & Bakh.f., Fl. Java 1 (1963) 495; Airy Shaw, Kew Bull. 26 (1972) 308; D.J.Rogers & Appan, Fl. Neotropica 13 (1973) 19; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 174; Kew Bull. Add. Ser. 8 (1980) 172; Kew Bull. 37 (1982) 30; G.L.Webster, Ann. Missouri Bot. Gard. 81 (1994) 99; Welzen, Nguyen & Chung, Rheedea 7 (1997) 79; Radcl.-Sm., Gen. Euphorbiacearum (2001) 273; Welzen, Q.D.Nguyen & R.K.C.Chung in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 437; G.L.Webster in Kubitzki, Fam. Gen. Vasc. Pl. 11 (2014) 162. — Janipha Humb., Bonpl. & Kunth, Nov. Gen. Sp. 2 (1817) 84 (folio), 106 (quarto), t. 109. — Mandioca Link, Handbuch 2 (19831) 436, nom. superfl. — Type: Jatropha manihot L. [= Manihot esculenta Crantz]
Manihotoides D.J.Rogers & Appan, Fl. Neotropica 13 (1973) 247. — Type: Manihotoides pauciflora (Brandegee) D.J.Rogers & Appan [= Manihot pauciflora Brandegee]
Monoecious subshrubs, shrubs, trees or vines. Roots tuberous and often with large quantities of stored starch. Stems with a soft wide pith, usually branching dichotomously, mainly sympodial; latex white, usually abundant. Indumentum simple hairs only. Stipules persistent to caducous, setaceous, margins entire, serrulate or dentate, leaving well-defined scars. Leaves alternate, simple; petioles long (SE Asia), often reddish; blade (SE Asia) palmatifid with (1–)5(–11 or more) lobes, seldom not lobed; lobes usually obovate, usually asymmetric except for the central lobe, papery, base (SE Asia) not to distinctly peltate, margin entire, glabrous (SE Asia), usually glaucous below; venation looped and closed near margin, usually scalariform. Inflorescences terminal or pseudo-axillary (due to development of side-branches, then opposite leaf) raceme-like panicles, usually several per node, branching sparingly once (SE Asia) with short branches; lower flowers pistillate, upper ones staminate, seldom flowers of one sexe only present; bracts and bracteoles subtending side-branches and flowers, margin entire, serrulate, serrate or laciniate. Flowers actinomorphic, 5-merous, pedicelled, abscission zone not visible; staminate flowers smaller than pistillate flowers; calyx in SE Asia campanulate, 5-lobed, lobes imbricate; petals absent. Staminate flowers: stamens 10, in 2 whorls of 5, outer larger (SE Asia), more or less outside disc glands, inner whorl among disc glands, filaments slender, anther versatile, subbasally dorsifixed, introrsely opening with longitudinal slits, 2-locular, connective elongated, elongation pilose; disc glands 10, radially arranged; pistillode small. Pistillate flowers: calyx lobes 5-nerved; staminodes sometimes present; disc annular, surface wobbly, glabrous; ovary 3-locular, one anatropous ovule per locule; style short, glabrous; stigmas 3, apically broadening, often split, glabrous, at the margin with globose stigmatic glands. Fruits capsular, globose to broadly ellipsoid, in Asia septicidal and partly, apically loculicidal; exocarp dry, smooth to warty, sometimes winged or ridged, often dehiscing from mesocarp; column hardly thickened basally and apically, with hardly any remains of the septa, subbasally with one vascular bundle, subapically 3 veins. Seed ellipsoid, dorsoventrally flattened, usually marbled with various shades of brown and white, shiny, apically with a caruncle. Embryo flat, surrounded by copious endosperm.
|
1a. |
Mainly small trees. Leaves distinctly (more than 5 mm) peltate. Ovary smooth. Fruits without wings. (Leaf lobes broad, lobes 0.9–3 times as long as wide; central unlobed part usually longer, lobes 5.8–12.5 times as long; nerves usually distinct and differently coloured when dry. Seeds 12 mm or longer.) |
|
|
1b. |
Mainly shrubs. Leaves not or slightly (up to 2 mm) peltate. Ovary with 6 longitudinal ridges; fruits with 6 longitudinal wings. (Leaf lobes narrow, c. 2.9–12.5 times as long as wide; central unlobed part usually short, lobes 15–21 times as long; nerves usually not very distinct and differently coloured when dry. Seeds up to 12 mm long. M. dulcis form in Malay Peninsula exceptional in these characters and like M. carthaginensis subsp. glaziovii). |
Manihot carthaginensis (Jacq.) Müll.Arg. subsp. glaziovii (Müll.Arg.) Allem, Novon 11 (2001) 160; Welzen, Q.D.Nguyen & R.K.C.Chung in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 438, Fig. 39: H, I. --- Manihot glaziovii Müll.Arg. in Mart., Fl. Bras. 11, 2 (1874) 446; Pax in Engl., Pflanzenr. IV.147.ii (1910) 89, fig. 31; Backer & Bakh.f., Fl. Java 1 (1963) 496; D.J.Rogers & Appan, Fl. Neotropica 13 (1973) 177; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 174; Kew Bull. Add. Ser. 8 (1980) 173; Welzen, Nguyen & Chung, Rheedea 7 (1997) 82. — Type: Glaziou 1022 (BR, G, n.v.; photo at F, n.v.), Brazil, near Rio de Janeiro.
(Shrubto) tree up to 13 m high, dbh up to 20 cm. Bark pinkish grey to dark brown, scaly, c. 0.6 cm thick, Araucaria-like; inner bark brown, thin; exudate thick, white latex; wood white. Stipules entire. Leaves: petiole green with reddish tinge; blade peltate, inserted 5–15(–20) mm from the margin; dull to shiny dark green above, green to pale bluish green below, latex present; lobes 0.9–3 times as long as wide; central unlobes part relatively long, lobes 5.8–12.5 times as long; nerves 7–17 pairs, venation very distinct, usually differently coloured when dry, veins scalariform. Inflorescences lax, usually single or few together. Staminate flowers: calyx devided to more or less halfway, green to greenish white to dark purple inside at base, glabrous; filaments white, anthers yellow. Pistillate flowers: calyx green tinged dull reddish to inside pinkish, glabrous; ovary smooth, yellow. Fruits globose, green with yellow sutures, warty, without wings. Seeds 12 mm or longer; caruncle yellow.
Distribution — Native to N Brazil. Sparsely cultivated throughout SE Asia and
Habitat & Ecology — Cultivated, often left behind and therefore found in scrubs, secondary forest and waste places along roads and rivers, near paddies, old cultivated fields; locally common. Often associated with Pterocarpus when growing along water. Soil: sticky red soil, may be water logged. Altitude: sea level up to 1200 m. Flowering whole year through, fruiting probably also, though no data available for August to December.
Uses — Cultivated all over Malesia for its latex which can be turned into rubber. However, Hevea brasiliensis outcompeted this species and presently it is hardly cultivated. The leaves can be eaten as vegetable, but must be chopped and cooked due to the HCN content of the leaves. The plant is often used on Java as a scion on Manihot esculenta roots to improve their production.
Vernacular names — English: Ceara rubber. Java: Oebi kajoe, oebi karet. Borneo: Kalimantan: Singkang tahunan; Sarawak: Bandong tujuh (tahun) (Selakau); ubi jipon, ubi tahun (Malay).
Note — See note 3 under M. esculenta.
Manihot esculenta Crantz, Inst. Rei Herb. 1 (1766) 167; Backer & Bakh.f., Fl. Java 1 (1963) 496; Airy Shaw, Kew Bull. 26 (1972) 308; D.J.Rogers & Appan, Fl. Neotropica 13 (1973) 25; Airy Shaw, Kew Bull. Add. Ser. 4 (1975) 174; Kew Bull. Add. Ser. 8 (1980) 172; Kew Bull. 37 (1982) 30; Veltkamp & G.H. de Bruijn in Flach & Rumawas, Plant Res. SE Asia 9 (1996) 107; Welzen, Nguyen & Chung, Rheedea 7 (1997) 80; in Welzen & Chayam., Fl. Thailand 8, 2 (2007) 438, Fig. 39: A-G. —Jatropha manihot L., Sp. Pl. (1753) 1007. — Janipha manihot Humb., Bonpl. & Kunth, Nov. Gen. Sp. 2 (1817) 84 (folio), 108 (quarto). — Manihot utilissima Pohl, Pl. Bras. Ic. Decr. 1 (1827) 32, t. 24, nom. superfl., see note 1; Müll.Arg. in DC., Prodr. 15, 2 (1866) 1064; Pax in Engl., Pflanzenr. IV.147.ii (1910) 67, fig. 24. — Mandioca utilissima (Pohl) Link, Handb. 2 (1831) 436. — Manihot manihot (L.) Cockerell, Bull. Torrey Club 19 (1892) 95, nom. illeg. — Lectotype (designated by Rogers & Appan 1973): Merian, Dissertatio generatione metamorphosibus insectorum Surinamensium (1726) Fig. 4 & Fig. 5.
Jatropha dulcis J.F.Gmel., Onom. Bot. 5 (1772-1778) 7. — Mandioca dulcis (J.F.Gmel.) Parodi, An. Soc. Ci. Argent. 4 (1877) 127. — Manihot dulcis (J.F.Gmel.) Pax in Engl., Pflanzenr. IV.147.ii (1910) 71. — Type not indicated, no lectotype known.
Jatropha mitis Rottb., Descript. Surinam. 21 (1776).
Jatropha stipulata Vellozo, Fl. Flum. Ic. 10 (1825) 5, t. 82. — Type: Vellozo s.n.
Manihot aipi Pohl, Pl. Bras. Ic. et Descr. 1 (1827) 29. — Manihot palmata (Vellozo) Müll.Arg. var. aipi (Pohl) Müll.Arg. in DC., Prodr. 15, 2 (1866) 1062. — Manihot dulcis (J.F.Gmel.) Pax in Engl., Pflanzenr. IV.147.ii (1910) 71. — Type: Pohl 3777 (iso in W).
Manihot aipi Pohl var. lanceolata Pohl, Pl. Bras. Ic. et Descr. 1 (1827) 31. — Type: Pohl 3778, p.p. (iso in W).
Manihot aipi Pohl var. latifolia Pohl, Pl. Bras. Ic. et Descr. 1 (1827) 31. — Type: Pohl 3776 (iso in W).
Manihot aipi Pohl var. lutescens Pohl, Pl. Bras. Ic. et Descr. 1 (1827) 31. — Type: Pohl 3780 (iso in W).
Manihot utilissima Pohl var. castellana Pohl, Pl. Bras. Ic. et Descr. 1 (1827) 34. — Type: Pohl 3778, p.p. (iso in W).
Manihot utilissima Pohl var. sutinga Pohl, Pl. Bras. Ic. et Descr. 1 (1827) 34. — Type: Pohl 3781 (iso in W).
Manihot flabellifolia Pohl, Pl. Bras. Ic. et Descr. 1 (1827) 35. — Jatropha flabellifolia (Pohl) Steud., Nomencl. ed. 2, 1 (1840) 799. — Manihot palmata (Vellozo) Müll.Arg. var. flabellifolia (Pohl) Müll.Arg. in DC., Prodr. 15, 2 (1866) 1062. — Manihot dulcis (J.F.Gmel.) Pax var. flabellifolia (Pohl) Pax in Engl., Pflanzenr. IV.147.ii (1910) 72. — Type: Pohl 1188 (iso in F, G, M, W).
Manihot digitiformis Pohl, Pl. Bras. Ic. et Descr. 1 (1827) 36, t. 27. — Jatropha digitiformis (Pohl) Steud., Nomencl. ed. 2, 1 (1840) 799. — Manihot palmata (Vellozo) Müll.Arg. var. digitiformis (Pohl) Müll.Arg. in DC., Prodr. 15, 2 (1866) 1063. — Syntypes: Pohl 1371 (F, W); Pohl 1709 (G, K).
Manihot diffusa Pohl, Pl. Bras. Ic. et Descr. 1 (1827) 55— Jatropha diffusa Steud., Nomencl. ed. 2, 1 (1840) 799. — Manihot palmata (Vellozo) Müll.Arg. var. diffusa (Pohl) Müll.Arg. in DC., Prodr. 15, 2 (1866) 1062. — Manihot dulcis (J.F.Gmel.) Pax var. diffusa (Pohl) Pax in Engl., Pflanzenr. IV.147.ii (1910) 71. — Type: Martius s.n. (iso in W).
Jatropha paniculata Ruiz & Pavon ex Pax in Engl., Pflanzenr. IV.147.ii (1910) 71.
Manihot loureirii Pohl, Pl. Bras. Ic. et Descr. 1 (1827) 55. — Jatropha loureirii (Pohl) Steud., Nomencl. ed. 2, 1 (1840) 799. — Type: Loureiro s.n.
Manihot edule A.Rich. in R. de la Sagra, Fl. Cub. ed. hisp. 3 (1853) 208.
Manihot melanobasis Müll.Arg., Linnaea 34 (1865) 206. — Syntypes: Schomburgk 426 (F, OXF, P, W); Schomburgk 426/694 (K)Schomburgk 694 (F, NY, P).
Manihot sprucei Pax in Engl., Pflanzenr. IV.147.ii (1910) 71. — Manihot esculenta Crantz var. sprucei (Pax) Lanjouw, Euphorb. Surinam 33 (1931). — Type: Spruce 825 (iso in BM, F, K, NY, OXF, P, W).
Manihot flexuosa Pax & K.Hoffm. in Engl., Pflanzenr. IV.147. (1924) 195. — Syntypes: Luetzelburg 9 (9a, 9b?) (F, M); Luetzelburg 10 (M, NY).
Not included are the infraspecific taxa of Ciferri (Archiv. Bot. Forli 18, 1942), because they do not refer to types.
(Herb to) shrub (to treelet), up to 7 m high, dbh up to 20 cm, single to few stems, sparingly branching; branchlets light green to tinged reddish, nodes reddish. Bark smooth, light brown to yellowish grey; inner bark cream-green; exudate thin, watery; wood soft, creamy straw. Stipules entire or somewhat split. Leaves: petiole light greenish to red; blade basally attached or slightly (up to 2 mm) peltate, dark green above, pale light greenish greyish underneath, sometimes variegated; lobes narrow, 2.9–12.5 times as long as wide; central unlobed part usually short, lobes 15–21 times as long; nerves 5-18 pairs, usually not very distinct, especially above, usually not differently coloured when dry, veins reticulate to scalariform. Inflorescences lax, with 3–5 together in fascicles; pedicels light green to red. Staminate flowers: calyx divided to halfway or more, green to white to lobes white to reddish with white median band inside to red purple, glabrous except for apex of calyx tube and inner side of segments finely hairy; filaments white, anthers yellow; disc yellow to light orange. Pistillate flowers: calyx green with red margin and midrib, hairy along the margin and on the midrib inside; disc pink; ovary with 6 longitudinal ridges, green (with pinkish stripes) to orange; pistil and stigmas white. Fruit subglobose, green (to light yellow, white, dark brown), rather smooth, with 6 longitudinal wings. Seeds up to 12 mm long.
Distribution — Probably originally cultivated in NE Brazil, nowadays only known from cultivation and found throughout SE Asia and Malesia.
Habitat & Ecology — Cultivated or found in waste places of mainly abandoned gardens and along roads, probably not escaped; locally common. Soil: clay, limestone, red loam, peaty soil, usually well-drained. Altitude: sea level up to 1700 m. Flowering and fruiting whole year through, though less so in September and October. See also Veltkamp & de Bruijn 1996.
Uses — The roots constitute one of the world’s largest stock crops for starch. It is mainly used for human consumption, less for animal consumption and (far less) for industrial purposes, though this may be very different per country. The roots can rarely be eaten fresh and are usually cooked, steamed, fried or roasted when fresh or after drying or fermenting. It is advisable to peel, grind or cut, and dry the roots in order to diminish the contents of cyanogenic glucosides (mainly linamarin). Only sweet cassava can be eaten fresh in small quantities, because the peel contains most HCN; the bitter ones have to be treated due to a much higher HCN content. The leaves are also eaten, they contain reasonable amounts of carotene and vitamin C, though they too have to be cut in pieces and they have to be cooked. Food poisoning hardly occurs, because most people know how to prepare the roots and leaves. See also Veltkamp & de Bruijn 1996.
Plants, especially with variegated leaves, are also used in horticulture.
Vernacular names — English: Brazilian arrowroot, cassava, tapioca. French: Manioc. Thailand: man-sampalang (general), man-samrong (Central), mam-mai (Peninsular). Malay Peninsula: Hubiq khan (Semelai); ubi belanda, ubi benggala, ubi kayu. Sumatra: Oebi, oebi kajoe, oebi paid. Java: Katela djendral, pohon, pohon abang, telo markan, telotengkor (Javanese); ketela pohon (M.); sampen (Sundanese); ubi dangdur, ubi pongger (Malay); singkong, singkot pahit, ubi kaju. Borneo: Sabah: Ubi kayu (Malay); ubi kayu puteh (Brunei); Sarawak: Bandung kapok (Bidayuh); jabang (Iban). Philippines: Balangay, kamoting-kahoy (Bisaya); balanghoy (Samar-Leyte Bisaya); kahoy, kahoy patata (Ifugao); kamoting-kahoy (Tagalog); kasaba (Ilokano); sang-laykayo (Tagbanua). Lesser Sunda Islands: Alor: Anti, batako hong kika, batako hong koeli, batako hong liking, batako kang. Moluccas: Halmahera: Inggris; Soela Sanana: Kasbi. New Guinea: Irian Jaya: Baundo taindi (Kuman); wateni (Kebar); Papua New Guinea: Hofa hagaya (Okapa); kawara (Taisora); cassava, mandioca, tapiagose, tapioc, timango heme.
Notes — 1. Rogers & Appan (1973) refer for Manihot utillissima Pohl to a type specimen. However, in Pohl’s description he refers to the Linnaean name Jatropha manihot, which makes it either a superfluous name when it really is a new species (like Rogers & Appan interpreted it), or it is superfluous because Pohl was not the first to create a new name under Manihot as Crantz did so already in 1766. The latter interpretation is favoured here.
2. Many cultivars of M. esculenta exist but no satisfactory classification of them exists.
3. Several specimens of the Malay Peninsula and, to a lesser extent, from New Guinea have broad lobes, a long central leaf part and very distinct, differently coloured venation. This form can be referred to as the ‘M. dulcis’ form. They strongly resemble M. carthaginensis subsp. glaziovii but they lack the peltation and they still possess ribs or wings along the fruits.
