LEMNACEAE [in review]
fu ping ke
Li Heng; Elias Landolt
Herbs, mostly perennial, minute, aquatic, floating or submersed, reduced to small green bodies called fronds corresponding partly to leaf and partly to stem. Roots absent or 121, filiform, arising from lower side of frond; root hairs absent. Fronds 1many cohering together, circular or oblong to lanceolate and flat or convex on lower side (gibbous), or globular to ovoid, 0.41.5 cm, with or without veins; daughter fronds successively formed from 1 basal cavity or 2 basal pouches detaching or remaining attached for many weeks. Node (point of origin of veins, roots and daughter fronds) situated c. 1/3 from base towards apex. Turions (in some species) formed under unfavourable conditions, frondlike, but smaller, compact, sinking to bottom. Flowers (interpreted as inflorescences by some authors) rare in most species, 1 (or 2), emerging from one of the pouches or from a cavity near median line on upper surface; sepals and petals absent; stamens 1 or 2; ovary 1, bottle-shaped, 17-ovuled, tapering into short style; stigma 1, funnel-shaped. Fruit indehiscent, opening by bursting.
Five genera and 38 species: worldwide in aquatic ecosystems; four genera and eight species in China.
Lemnaceae species grow often together with other species of the same family, sometimes also with water ferns. They are usually gregarious and multiply rapidly (up to doubling within 24 hours). The fronds of all species are used as high quality feed for domestic animals, fish and fowl.
1a. Frond rootless, without veins; each frond with 1 basal cavity reproducing daughter fronds; flower emerging from a cavity near the median line of the upper surface, not surrounded by scale; stamen 1, 2-locular; seed 1, nearly smooth 4. Wolffia
1b. Frond with 121 roots, with 121 veins; each frond with 2 lateral pouches at the base reproducing daughter fronds and (rarely in most species) 1 flower; flower surrounded by small utricular, membranous scale; stamens 2, 4-locular; seeds 15, longitudinally ribbed.
2a. Frond with 1 root, with 15(7) veins, without a scale at base 3. Lemna
2b. Frond with 221 roots and (3)521 veins, surrounded at base by a small membranous scale covering the point of attachment of the roots.
3a. Frond 11.5 times as long as wide, with 716(21) veins, with (5)721 roots, flat or (rarely) slightly gibbous; 1 (or 2) roots perforate covering scale 1. Spirodela
3b. Frond 1.53 times as long as wide, with (3)57 veins, with (1)27(12) roots, flat or gibbous; all roots perforate covering scale 2. Landoltia
1. SPIRODELA Schleiden, Linnaea 13: 391. 1839
zi ping shu
Fronds free floating, 110 cohering together, usually asymmetrical, flat or sometimes convex (gibbous) on lower side, shiny green above, often red beneath, with 2 lateral pouches at base, reproducing daughter fronds and (rarely) 1 flower, surrounded at base by small membraneous scale covering point of attachment of the roots; veins (5)721, leaving from node towards apex, sometimes visible on upper surface. Pigment cells present (visible in dead fronds as brown dots). Roots 521, surrounded at base by a tubular sheath, enclosed at tip by a rounded to pointed cap. Daughter fronds connected to mother frond by thin white stipe. Flower surrounded by small utricular, membranous scale with narrow opening at top; stamens 2, 4-locular. Seeds 13, longitudinally ribbed.
Two species: one species worldwide, one species only in Central and South America; one species in China.
1. Spirodela polyrrhiza (Linnaeus) Schleiden, Linnaea 13: 392. 1839
zi ping
Lemna polyrhiza Linnaeus, Sp. Pl.: 970. 1753
Fronds obovate to circular, 310 mm, 11.5 times as long as wide, flat or (rarely) gibbous, sometimes with indistinct papillae on upper surface along veins. Roots (5)721, 1 (or 2) perforating scale, 0.53 cm. Turions sometimes present, without roots, circular to reniform, 12 mm, brownish to olive. Flower with 1 or 2 ovules. Fruit laterally winged towards top; seed with 1220 ribs. Fl. (very rare) JunSep. 2n = 30, 32, 38, 40*, 50, 80.
Ponds, lakesides, rice fields, pools, ditches; lacking in regions with cool summers. 02900 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hebei, Heilongjiang, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Jilin, Liaoning, Quinghai, Shaanxi, Shandong, Shanxi, Sichuan, Taiwan, Yunnan, Zhejiang [all continents].
Gibbous fronds of S. polyrrhiza are sometimes named var. masonii Daubs. This variant occurs rarely throughout the whole distribution and is of no systematic value.
S. polyrrhiza is used as Chinese medicinal herb for diuretics, for subsides of swellings and for reducing perspiration.
2. LANDOLTIA Les & Crawford, Novon 9: 532. 1999
see note at top
Fronds free floating, 110 cohering together, usually asymetrical, flat or gibbous, shiny green above, often red beneath, with 2 lateral pouches at base, reproducing daughter fronds and (rarely) 1 flower, surrounded at base by small membraneous scale covering point of attachment of the roots; veins (3)57, leaving from node towards apex, often visible on upper surface. Pigment cells present (visible in dead fronds as brown dots). Roots (1)27(12), surrounded at base by a tubular sheath, enclosed at tip by a rounded or pointed cap. Daughter fronds connected to mother frond by thin white stipe. Flower surrounded by small utricular, membraneous scale with a slit on one side (rarely on 2 sides); stamens 2, 4-locular. Seeds 1(2), longitudinally ribbed.
One species: originally probably distributed only in SE Asia and Australia, now introduced in subtropical and tropical zones worldwide; one species in China.
1. Landoltia punctata (G. Meyer) Les & Crawford, Novon 9: 532. 1999
see note at top
Lemna punctata G. Meyer, Prim. Fl. Esseq.: 262. 1818; Spirodela punctata (G. Meyer) C.H. Thompson; Spirodela oligorrhiza (Kurz) Hegelm.
Fronds obovate to elliptic, 1.58 mm, 1.53 times as long as wide, flat or gibbous, mostly with papillae on upper surface along midvein. Roots (1)27(12), all perforating scale, 0.57 cm. No distinct turions present. Flower with 1 or 2 ovules. Fruit laterally winged towards top; seed with 1015 distinct ribs. Fl. (rare) JunSep. 2n = 40*, 46, 50.
Ponds, lakesides, pools, ditches; in regions with mild winters and not very hot summers. 02400 m. Fujian, Henan, Hubei, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [India, Indonesia, Japan, Malaysia, Philippines, Thailand, Vietnam; Africa, Australia, North America, Pacific Islands, South America].
M.G. Liu and K.M. Xie described 1983 a new species of Spirodela from Sichuan under the name S. sichuanensis. The species has been distinguished from S. punctata (now Landoltia punctata) mainly by the form of the utricular scale surroundig the flower: the scale has 2 lateral slits in S. sichuanensis and one lateral slit in L. punctata. However, the 2 slits can rarely also be observed in other places of the distribution area of L. punctata. Due to the rarity of flowering, the frequency of this character is not known. All the vegetative characters of S. sichuanensis stay within the morphologiocal variation of L. punctata.
3. LEMNA Linnaeus, Sp. Pl. 2: 970. 1753
fu ping shu
Fronds free floating on the water surface or submersed, 1 to many cohering together, symmetrical or asymmetrical, flat or sometimes convex beneath (gibbous), green above, sometimes red beneath, with 2 lateral pouches at base, reproducing daughter fronds and (rarely in most species) 1 flower, not surrounded by a scale at base; veins 15(7), leaving from node towards apex. Pigment cells absent. Root 1, sometimes dropped off or lacking (L. trisulca), surrounded at base by a short tubular sheath and enclosed at tip by a cap. Daughter fronds connected to mother frond by a thin white stipe or a green stalk (L. trisulca). Flower surrounded by small utricular, membranous scale; stamens 2, 4-locular. Seeds 15, longitudinally ribbed.
Thirteen species: nearly cosmopolitan; five species in China.
1a. Fronds submersed (except when flowering or fruiting), often forming branched chains, at base narrowed into green stalk, margins denticulate distally 1. L. trisulca
1b. Fronds floating upon the water surface, singular or forming small groups, at base rounded, no green stalk present, margins entire.
2a. Frond with 1 vein 6. L. minuta
2b. Frond with 35 veins.
3a. Root sheath winged at base; root shorter than 3 cm; fronds without reddish colour 5. L. aequinoctialis
3b. Root sheath not winged at base; root 0.515(19) cm long; fronds sometimes slightly or deep red coloured or with reddish spots on lower or upper surface.
4a. Small, olivebrown, rootless turions often present which, sink to bottom 2. L. turionifera
4b. Plants without distinct turions.
5a. Fronds with mostly distinct papillae on midline of upper surface; greatest distance between lateral veins near or distal to middle 2. L. turionifera
5b. Fronds with distinct papillae only above node and apex; greatest distance between lateral veins near or proximal to middle.
6a. Fronds often reddish or red on lower surface (more intensely so than on upper), flat or slightly gibbous 3. L. japonica
6b. Fronds always green on lower side, sometimes reddish on upper surface, flat 4. L. minor
1. Lemna trisulca Linnaeus, Sp. Pl. 2: 970. 1753.
pin zao
Fronds (except when flowering or fruiting) submersed, 350, coherent in groups and often forming branched chains, transparent green, sometimes purplish, narrowed at base into a 220 mm long green stalk, 315 mm (without stalk), 23.5 times as long as wide, narrowly ovate to elliptic, flat, without papillae on upper surface; margins denticulate distally; veins (1 or) 3, central vein reaching nearly apex, lateral veins only about halfway between node and apex. Root 0.52.5 cm, often lacking; sheath not winged; tip mostly pointed. Turions absent. Flowering and fruiting fronds free floating on water surface, 15, coherent, 35 mm. Flower with 1 ovule; utricular scale with narrow opening at top. Fruit laterally winged towards top; seed with 1218 distinct ribs. Fl. (rare) MaySep. 2n = 20, 40, 42, 60, 63, 80.
Lakesides, spring pools, ponds; in regions with cool temperate climate. 03000 m. Anhui, Hebei, Heilongjiang, Hubei, Jiangsu, Nei Mongol, Shaanxi, Shanxi, Sichuan, Taiwan, Xinjiang, Yunnan, Zhejiang [Afghanistan, India, Japan, New Guinea, Kasakhstan, Kyrgyzstan, Pakistan, Philippines, Russia; all continents except South America].
2. Lemna turionifera Landolt, Aquatic Botany 1: 355. 1975.
name
Fronds free floating on water surface, 18, coherent in small groups, shiny green, sometimes with reddish spots on upper surface and red coloured on lower surface (always deeper red on lower surface than on upper), rounded at base, without a green stack (but with thin white stipe connecting fronds), 1.54 mm, 11.5 times as long as wide, obovate, flat, with distinct papillae on midline of upper surface; margins entire; veins 3, reaching nearly apex, greatest distance between lateral veins near or distal to middle. Root 0.515 cm; sheath not winged, tip rounded (to pointed). Turions often present, 0.51.5 mm, olivebrown, rootless. Flower with 1 ovule; utricular scale with narrow opening at top. Fruit not winged towards top; seed with 3060 indistinct ribs. Fl. (rare) JunSep. 2n = 40, 42, 50, 80.
Lakesides, spring pools, ponds, slowly flowing streems; in regions with continental cool temperate climate. 03000 m. Anhui, Hebei, Heilongjiang, Nei Mongol [Korea, N Japan, Russia; North America, Asia, Europe (introduced)].
3. Lemna japonica Landolt, Veröff. Geobot. Inst. ETH, Stiftg. Rübel, 70: 23. 1980.
name
Lemna minor auct mult. non Linnaeus
Fronds free floating on water surface, 18, coherent in small groups, shiny green, sometimes reddish or red on lower surface (always deeper red on lower surface than on upper), rounded at base, without a green stalk (but with thin white stipe connecting fronds), 26(7) mm, 1.31.8 times as long as wide, obovate to elliptic, flat or sligtly convex on lower side, with mostly distinct papillae on upper surface near apex and above node and indistinct ones along the midline; margins entire; veins 3(5), reaching nearly apex, greatest distance between lateral veins near or proximal to middle. Root 0.515(18) cm; sheath not winged, tip rounded (to pointed). Turions absent. Flower with 1 ovule; utricular scale with narrow opening at top. Fruit not known. Fl. (very rare) JulOct. 2n = 40*, 50, 63.
Lakesides, pools, ponds, ditches; in regions with moderate to warm temperate climate. 02900 m. Hebei, Heilongjiang, Henan, Hubei, Jiangsu, Nei Mongol, Shaanxi, Shandong, Shanxi, Yunnan, Zheijiang [Korea, Japan].
L. japonica probably originated from hybridization between L. minor and L. turionifera.
M.G. Liu & C.H. You described 1983 Lemna leibonensis from S. Sichuan which is most probably identical with L. japonica. It is distinguished by larger fronds, conspicious veins and pointed root caps. These characters occur also within L. japonica..
4. Lemna minor Linnaeus, Sp. Pl. 2: 970. 1753.
fu ping
Fronds free floating on water surface, 18, coherent in small groups, shiny green, sometimes reddish on upper surface, scarcely reddish on lower surface (always deeper red on upper surface than on lower), rounded at base, without a green stalk (but with thin white stipe connecting fronds), 26(10) mm, 1.32 times as long as wide, obovate to elliptic, flat, with mostly distinct papillae on upper surface near apex and above node; margins entire; veins 3(5), reaching nearly apex, greatest distance between lateral veins near or proximal to middle. Root 0.515 cm; sheath not winged, tip mostly rounded. Turions absent. Flower with 1 ovule; utricular scale with narrow opening at top. Fruit winged laterally towards top; seed with 1016 distinct ribs. Fl. (rare) MaySep. 2n = 20, 40, 42, 50, 63, 126.
Lakesides, pools, ponds, ditches, slow flowing streams in regions with suboceanic cool to moderate temperate climate. (0) 20003000 m. W. Xizang [Afghanistan, N India, Kazakhstan, Nepal, N Pakistan, W Russia, Turkmenistan; Africa, SW Asia, Japan (introduced), Europe, North America, Australia (introduced), New Zealand (introduced)].
5. Lemna aequinoctialis Welwitsch, Apont. Conselh. Ultramar. 55: 578. 1859.
xi mai fu ping
Lemna paucicostata Hegelmaier; Lemna perpusilla auct. mult. non Torrey; Lemna trinervis (Austin) Small
Fronds free floating on water surface, 18 coherent in small groups, shiny bright green, without reddish colour, rounded at base, without a green stalk (but with thin white stipe connecting fronds), 1.54(6.5) mm, 13 times as long as wide, obovate to lanceolate, flat, with mostly very distinct papillae on upper surface near apex and above node; margins entire; veins 3, reaching nearly apex, greatest distance between lateral veins nearly or proximal to middle. Root 0.53 cm; sheath winged towards the base, tip mostly pointed. Turions absent. Flower with 1 ovule; utricular scale with slit on one side. Fruit not winged; seed with 824 distinct longitudinal ribs. Fl. (frequent) all seasons. 2n = 20, 40*, 42, 50*, 60, 70, 80, 84.
Lakesides, pools, ponds, rice fields, ditches; in regions with warm temperate to tropical climate. 02800 m. Anhui, Fujian, Guangdong, Guizhou, Hebei, Henan, Hubei, Jiangsu, Jiangxi, Liaoning, Quinghai, Shaanxi, Shandong, Shanxi, Taiwan, Yunnan, Zhejiang [Afghanistan, Bangladesh, Bhutan, Cambodia, India, Indonesia, Japan, Korea, Laos, Malaysia, Myanmar, Nepal, New Guinea, Pakistan, Philippines, Singapore, Sri Lanka, Thailand, Vietnam; all continents].
Plants with large thin fronds with conspicuous veins and without distinct papillae are often called L.trinervis. This character combination represents just a modification occurring throughout the range of L. aequinoctialis.
6. Lemna minuta Kunth in A. von Humboldt & al., Nov. Gen. Sp. 1: 372. 1816.
name
Lemna minima Phil.; Lemna minuscula Herter
Fronds free floating on water surface, 18, coherent in small groups, bright green, without reddish colour, rounded at base, without a green stalk (but with thin white stipe connecting fronds), 0.84 mm, 12 times as long as wide, obovate, flat to thickish (but not convex on lower side), with or without indistinct papillae on midline of upper surface; margin entire; vein 1, not longer than 2/3 of the distance between node and apex. Root 0.21.5 cm; sheath not winged, tip rounded to pointed. Turions absent. Flower with 1 ovule; utricular scale with slit on one side. Fruit not winged; seed with 1215 distinct ribbs. Fl. (very rare) MaySep. 2n = 36, 40, 42.
Lakesides, spring pools, ponds, ditches; in regions with suboceanic moderate to warm temperate climate. 0-->3000m. Not known from China but likely to be introduced [C Japan (introduced); America, Europe (introduced)].
4. WOLFFIA Horkel ex Schleiden, Beitr. Bot.: 233. 1844.
wu gen ping shu
Fronds free floating on or below the water surface, 1 or 2 cohering together, symmetrical, globular, ovoid or boat-shaped, green above, green to transparent beneath, never reddish, with 1 basal cavity reproducing daughter fronds, not surrounded by a scale at base; veins absent. Pigment cells (in our species) absent in vegetative cells. Roots absent. Daughter fronds connected to mother frond by a very short stipe (not visible). Flower emerging from a cavity near median line of upper surface, not surrounded by a scale; stamens 1, 2-locular. Seed 1, nearly smooth.
Eleven species: nearly cosmopolitan; one species in China.
1. Wolffia globosa (Roxburgh) Hartog & Plas, Blumea 18: 367. 1970.
wu gen ping
Lemna globosa Roxburgh, Fl. Ind. 3: 565. 1832;Wolffia arrhiza auct. mult. non (Linnaeus) Horkel
Fronds free floating on or just below water surface, green on upper surface, transparent green below, ovoid, 0.40.8 mm, 1.32 times as long as wide, 11.5 times as deep as wide, rounded or sligtly pointed at apex; papillae absent. Turions sometimes present, 0.30.5 mm, whitish green, globular. Fl. (very rare) JunSep. 2n = 16, 23, 30, 40*, 46, 50, 60.
Lakesides, pools, ponds and ditches; in regions with suboceanic, warm temperatetropical climate. 01300 m. Fujian, Guangdong, Henan, Hubei, Jiangsu, Taiwan, Yunnan, Zhejiang [Bangladesh, Cambodia, India, Indonesia, Japan, Laos, Malaysia, Myanmar, Nepal, Pakistan, Philippines, Singapore, Sri Lanka, Thailand, Vietnam; introduced to America].