SPARGANIACEAE [Draft]

黑三棱科  hei san leng ke

Sun Kun (孙坤)[1]; David A. Simpson[2]

Herbs, perennial, aquatic or of marshes, with creeping rhizomes. Stems simple or branched, robust or slender. Leaves alternate, distichous, emergent or floating, sheathed at base. Plants monoecious; flowers minute, densely crowded in globose heads on upper part of stems and branches; several staminate and pistillate heads in panicle, raceme, or spike; staminate heads above pistillate. Perianth segments of 3–6 chaffy elongate scales. Staminate flowers with 3 or more stamens; anthers basifixed, longitudinally dehiscent; filaments free or partly united. Pistillate flowers bracteolate; ovary 1-loculed or rarely 2-loculed, narrow at base; styles simple or forked, with a unilateral stigma; ovule 1, pendulous. Fruit nutlike, indehiscent. Seeds with thin testae.

One genus and about 19 species: mainly in temperate and cool regions of the world, best represented in the northern hemisphere; 11 species (three endemic) in China.

1. SPARGANIUM Linnaeus, Sp. Pl. 2: 971. 1753.

黑三棱属  hei san leng shu

Morphological characters and geographical distribution are the same as those of the family.

??Sparganium affine Schnizl.

??S. asiaticum Graebn.

??S. carinatum Falc.

??S. coreanum - see Fl. Liaoning

??S. emersum - see Fl. Liaoning

??S. fluitans Fr.

??S. glehnii Meinsh.

??S. japonicum Rothert

??S. longifolium Turcz.

??S. microcarpum celax - see Fl. Xinjiang

??S. polyedrum Asch. & Graebn.

??S. ramosum subsp. neglectum Beeby, J. Bot. 23: 26, 193, t. 285. 1885.

1a.       Stems floating or slightly prostate in water; leaves usually floating, flattened.

2a.       Stems stout, 1.3–1.5 m tall; leaves ca. 10 mm wide; staminate heads 8–10
............................................................................................................  8. S. yunnanense

2b.       Stems slender, less than 70 cm tall; leaves 2–4 mm wide; staminate heads 1–4.

3a.       Styles absent or inconspicuous; fruit broadly obovoid .....................  11. S. hyperboreum

3b.       Styles conspicuous; fruit fusiform, elliptical, or lanceolate.

4a.       Stems floating, more than 20 cm; staminate heads 2–4 ..................  9. S. angustifolium

4b.       Stems slightly prostate or rarely erect, less than 20 cm; staminate heads 1 or 2  10. S. minimum

1b.       Stems erect; leaves aerial, not flattened.

5a.       Inflorescences paniculate, with conspicuous lateral branches with staminate and pistillate heads; ovary sessile.

6a.       Panicles with 3–7 lateral branches; stigmas 3–4 mm ..........................  1. S. stoloniferum

6b.       Panicles with 1(or 2) lateral branches; stigmas less than 2 mm.

7a.       Leaves 2–3 mm wide; main axis and lateral branch of panicles not zigzag; styles ca. 0.5 mm             2. S. stenophyllum

7b.       Leaves 4–5 mm wide; main axis and lateral branch of panicles usually zigzag; styles 1–1.5 mm         3. S. limosum

5b.       Inflorescences racemose or spicate, degenerated into one pistillate head or completely degenerated; ovary with short stalk.

8a.       Inflorescences spicate; perianth segments lobate to parted .....................  7. S. confertum

8b.       Inflorescences racemose; perianth segments serrate-lobate to lobate.

9a.       Axis of racemes curved; ovary attenuate toward base and forming an inconspicuous stalk  4. S. fallax

9b.       Axis of racemes not curved; ovary with a conspicuous stalk.

10a.     Staminate heads 4–8, distinctly separate from pistillate ones; stigmas 1.5–1.8 mm  5. S. simplex

10b.     Staminate heads 1–3, not separate from pistillate ones; stigmas ca. 0.5 mm  6. S. glomeratum

1. Sparganium stoloniferum (Graebner) Buchanan-Hamilton ex Juzepczuk in Komarov, Fl. URSS 1: 219. 1934.

黑三棱  hei san leng

Stems emersed, erect, robust, 70–120 cm tall or more. Leaves (20–)40–90 Χ 0.7–1.6 cm, upper flattened, lower keeled or trigonous on abaxial side. Panicles (8–)20–60 cm, with 3–7 lateral branches, each with (1 or)7–11 staminate heads and 1 or 2 pistillate heads; main axis consisting of 3–5 or more staminate heads. Male flowers: perianth segments spatulate, lobate at apex; filaments ca. 3 mm; anthers obconical, (0.5–)1–1.2 Χ ca. 0.5 mm. Female flowers: perianth segments 5–7 Χ 1–1.5 mm; ovary sessile; styles forked or simple, ca. 1.5 mm; stigmas 3–4 mm. Fruit obconical, (3.5–)6–9 mm, ribbed. Fl. and fr. May-Oct.

In lakes, streams, swamps, and ponds, up to ca. 3600 m. ??Anhui, Gansu, Hebei, Heilongjiang, ??Henan, Hubei, Jiangsu, Jiangxi, Jilin, Liaoning, Nei Mongol, Shaanxi, ??Shandong, Shanxi, Xinjiang, Xizang, Yunnan, ??Zhejiang [Afghanistan, Japan, Korea, ??Mongolia, Russia; C Asia – list countries adjacent to China??].

1a. Panicles 20–60 cm, each branch with 7–11 staminate heads; anthers 1–1.2 mm; fruit 6–9 mm ..  1a. subsp. stoloniferum

1b. Panicles 8–15 cm, each branch with only 1 staminate head; anthers ca. 0.5 mm; fruit ca. 3.5 mm  1b. subsp. choui

1a. Sparganium stoloniferum subsp. stoloniferum

黑三棱(原亚种)  hei san leng (yuan ya zhong)

Sparganium ramosum Hudson subsp. stoloniferum Graebner in Engler, Pflanzenr. 2(IV.10): 14. 1900; S. arcuscaulis D. Yu & G. T. Yang; S. multiporcatum D. Yu.

Panicles 20–60 cm, each branch with 7–11 staminate heads. Anthers 1–1.2 mm. Fruit 6–9 mm.

Lakes, ponds, swamps, streams; below 3600 m. Gansu, Hebei, Heilongjiang, Hubei, Jiangsu, Jiangxi, Jilin, Liaoning, Nei Mongol, Shaanxi, Shanxi, Xinjiang, Xizang, Yunnan [Afghanistan, Japan, Korea, Russia; C Asia].

Some specimens from Heilongjiang were respectively described as Sparganium multiporcatum D. Yu (Bull. Bot. Res., Harbin 11(2): 20. 1991) and S. arcuscaulis D. Yu & G. T. Yang (loc. cit.: 37), distinguished either for the female flowers with perianth segments slightly lobate, stigmas 1.5–2 mm, or for the stems arcuate at the upper part, fruit slightly small. They represent local variant individuals rather than distinct taxa.

1b. Sparganium stoloniferum subsp. choui (D. Yu) K. Sun, comb. nov.

周氏黑三棱  zhou shi hei san leng

Basionym: Sparganium choui D. Yu, Bull. Bot. Res., Harbin 12: 257. 1992.

Panicles 8–15 cm, each branch with only one staminate head; anthers ca. 0.5 mm; fruit ca. 3.5 mm.

* Nei Mongol.

2. Sparganium stenophyllum Maximowicz ex Meinshausen, Bull. Soc. Imp. Naturalistes Moscou, n.s., 3: 171. 1889.

狭叶黑三棱  xia ye hei san leng

Stems erect, 20–36 cm tall, slender. Leaves 25–35 cm Χ 2–3 mm, upper flattened, lower keeled or trigonous on abaxial side. Panicles 7–15 cm, main axis consisting of 5–7 staminate heads and 2 or 3 pistillate heads, usually with 1 lateral branch at base; lateral branch 5–8 cm, with 2 or 3 staminate and 1 or 2 pistillate heads. Male flowers: perianth segments spatulate, ca. 2 mm, lobate at apex; filaments ca. 2 mm; anthers oblong, ca. 1 Χ 0.3 mm. Female flowers: perianth segments spatulate, ca. 2 mm, lobate at apex; ovary sessile; styles ca. 0.5 mm; stigmas unilateral, ca. 1.5 mm. Fruit obovoid, ca. 4 mm. Fl. and fr. Jun–Sep.

Shallow water of lakes, swamps, and streams. Hebei, Heilongjiang, Jilin, Liaoning [Japan, Korea, Russia].

3. Sparganium limosum Y. D. Chen, Acta Phytotax. Sin. 19: 52. 1981.

沼生黑三棱  zhao sheng hei san leng

Stems emersed, erect, 33–57 cm tall. Leaves erect, 50–70 cm Χ 4–5 mm, upper flattened, lower keeled or trigonous on abaxial side. Panicles 7–16 cm, with 1 lateral branch at base, main axis and lateral branch usually zigzag; main axis with 5–8 staminate heads and 2 or 3 pistillate heads; lateral branch with 3(–5) staminate and 1 pistillate heads. Male flowers: perianth segments ca. 1.5 Χ 0.2 mm; filaments 3–4 mm; anthers oblong, ca. 1 Χ 0.2 mm. Female flowers: perianth segments 1.7–2.5 Χ ca. 1 mm, serrate-lobate at apex; ovary sessile; styles 1–1.5 mm; stigmas linear, 1–1.2 mm. Fl. and fr. Jun–Sep.

* Swamps; ca. 1800 m. NW Yunnan (Gaoligong Shan).

4. Sparganium fallax Graebner, Allg. Bot. Z. Syst. 4: 32. 1898.

曲轴黑三棱  qu zhou hei san leng

Stems emersed, erect, 40–55(–80) cm tall. Leaves 45–65 cm Χ 4–10 mm, upper flattened, lower keeled or trigonous on abaxial side. Racemes 15–17 cm, basally curved; staminate heads 4–7, distinctly separated from pistillate ones; pistillate heads 3 or 4(–7), relatively widely separated, sessile or lowest pedunculate. Male flowers: perianth segments linear, 2–2.5 mm, serrate-lobate at apex; filaments 3–4.5 mm; anthers oblong, ca. 1.5 Χ 0.4 mm. Female flowers: perianth segments broadly spatulate, ca. 3 Χ 1–1.5 mm, serrate-lobate at apex; ovary attenuate toward base; styles short; stigmas 1.5–2 mm. Fruit broadly fusiform, 5–6 mm. Fl. and fr. Jun-Oct.

Shallow water of lakes, ponds, swamps, and streams. Fujian, Guizhou, Taiwan, Yunnan, Zhejiang [India, Japan, Myanmar].

5. Sparganium simplex Hudson, Fl. Angl., ed. 2, 401. 1778.

小黑三棱  xiao hei san leng

Stems emersed, erect, 30–70 cm tall, slender. Leaves 40–80 cm, aerial or floating, lower keeled on abaxial side. Racemes 10–20 cm; staminate heads 4–8, distinctly separate from pistillate ones; pistillate heads 3 or 4, relatively widely separated, sessile or lowest 1 or 2 pedunculate. Male flowers: perianth segments linear or spatulate, 2–2.5 mm, lobate at apex; filaments ca. 4 mm; anthers oblong, ca. 1.6 Χ 0.4 mm. Female flowers: perianth segments spatulate, ca. 3.5 mm, lobate at apex; ovary with short stalk; styles ca. 1 mm; stigmas 1.5–1.8 mm. Fruit fusiform. Fl. and fr. Jun–Oct. 2n = 30.

Margins of lakes, swamps, and streams. Gansu, ??Hebei, Heilongjiang, ??Henan, Jilin, Liaoning, Nei Mongol, ??Shaanxi, Xinjiang [Japan, ??Myanmar, Russia; elsewhere??].

6. Sparganium glomeratum Laestadius ex Beurling, Φfvers. Fφrh. Kongl. Svenska Vetensk.-Akad. 9: 192. 1853.

短序黑三棱  duan xu hei san leng

Sparganium manshuricum D. Yu.

Stems emersed, erect, 20–50 cm tall. Leaves 30–56 cm, lower keeled or trigonous on abaxial side. Racemes 6–15 cm; staminate heads 1–3, not separate from pistillate ones; pistillate heads 3 or 4(–8), relatively near each other, lowest one pedunculate. Male flowers: perianth segments ca. 1.5 mm, serrate-lobate at apex; filaments 3–3.5 mm; anthers oblong, ca. 1 mm. Female flowers: perianth segments 2–2.5 mm, lobate at apex; ovary fusiform, with short stalk; styles short; stigmas ca. 0.5 mm. Fruit broadly fusiform, ca. 3 mm. 2n = 30. Fl. and fr. Jun–Sep.

Margins of lakes, swamps, and slow-moving rivers. Heilongjiang, Jilin, Liaoning, Nei Mongol, Xizang, Yunnan [Japan, ??Mongolia, Russia; Europe].

A specimen with 7 or 8 pistillate heads and larger flowers from Heilongjiang, which was described as Sparganium manshuricum by D. Yu (Bull. Bot. Res., Harbin 12: 255. 1992) mainly according to some quantitative characters, perhaps represents a local variant suite of specimens rather than a distinct taxon.

7. Sparganium confertum Y. D. Chen, Acta Phytotax. Sin. 19: 53. 1981.

穗状黑三棱  sui zhuang hei san leng

Stems emersed, erect, 15–45 cm tall. Leaves 25–55 cm Χ 3–5 mm, lower keeled on abaxial side. Spikes with many densely arranged heads, 6–11 cm; staminate heads (6–)8–12, deciduous; pistillate heads 1–3, sessile or rarely lowest one sterile, pedunculate. Female flowers: perianth segments broadly ??, 3–5 Χ ca. 1.5 mm, lobate to parted at apex; ovary elliptical, with stalk 1.5–2 mm; styles ca. 1 mm; stigmas unilateral, 1–1.5 mm. Fruit 4–5 mm. Fl. and fr. Sep–Oct.

* Shallow water of rivers, swamps, and ponds; ca. 3100 m. NW Yunnan (Gaoligong Shan).

8. Sparganium yunnanense Y. D. Chen, Acta Phytotax. Sin. 19: 54. 1981.

云南黑三棱  yun nan hei san leng

Stems floating, 1.3–1.5 m long, stout. Leaves floating, ca. 1.5 m Χ ca. 1 cm, flattened, spongy, lower slightly convex on abaxial side. Racemes 20–35 cm, lower curved?; staminate heads 8–10; pistillate heads 3 or 4(–7), sessile or lowest 1 or 2 pedunculate. Male flowers: perianth segments spatulate, 3–3.5 mm; filaments 3–4 mm; anthers linear, ca. 2 mm. Female flowers: perianth segments white, spatulate, 4–5 mm, lobate at apex; ovary fusiform, attenuate toward base; styles ca. 1.5 mm; stigmas oblong to ovate, 1.5–2 mm, lacerous or forked. Fl. and fr. May–Sep.

* About 1500 m. SW Yunnan (Menghai).

9. Sparganium angustifolium Michaux, Fl. Bor.-Amer. 2: 189. 1803.

线叶黑三棱  xian ye hei san leng

Stems floating, 30–50 cm long, slender. Leaves floating, 25–40 cm Χ ca. 4 mm, flattened. Racemes 6–16 cm; staminate heads 2–4; pistillate heads 2–4, relatively widely separated, sessile or lowest 1 or 2 pedunculate. Male flowers: perianth segments spatulate to obtriangular; filaments ca. 5 mm; anthers elliptical, 1–1.2 Χ ca. 0.5 mm. Female flowers: perianth segments linear to spatulate, serrate-lobate to parted; ovary fusiform, attenuate toward base and forming a short stalk; styles ca. 2 mm; stigmas narrowly elliptical. Fruit fusiform to elliptical, ca. 4 mm. Fl. and fr. Jul–Sep. 2n = 30.

Lakes, streams, swamps; above 1500 m. Heilongjiang, Jilin, Xinjiang [Japan [elsewhere in N Asia??]; Europe, North America].

10. Sparganium minimum Wallroth, Erst. Beitr. Fl. Hercyn. 2: 297. 1840.

矮黑三棱  ai hei san leng

Stems slightly prostate or rarely erect, 1.5–20 cm long, slender. Leaves aerial or floating, 20–25 cm Χ ca. 4 mm, flattened. Spikes or racemes 2–3.5 cm; staminate heads 1(or 2); pistillate heads 2(or 3), lowest one sometimes pedunculate. Male flowers: perianth segments ca. 1.5 mm, lobate at apex; filaments ca. 2 mm; anthers oblong, ca. 1 mm. Female flowers: perianth segments spatulate, ca. 2 mm, lobate to parted at apex; ovary lanceolate, with or without short stalk; styles short; stigmas ca. 1 mm. Fruit broadly lanceolate. Fl. and fr. Jul–Sep. 2n = 30.

Below 3500 m. Heilongjiang, Nei Mongol, Sichuan [??Mongolia, Russia; Europe, North America].

Specimens described as Sparganium amplexicaulium D. Yu (Bull. Bot. Res., Harbin 12: 257. 1992) and S. tenuicaule D. Yu & L. H. Liu (Bull. Bot. Res., Harbin 11(2): 19. 1991) are similar to this species. Because the two species were mainly described according some quantitative characters and based on only one specimen, respectively, their taxonomic positions need to be further studied.

11. Sparganium hyperboreum Laestadius ex Beurling, Φfvers. Fφrh. Kongl. Svenska Vetensk.-Akad. 9: 192. 1853.

无柱黑三棱  ?? ?? hei san leng

Stems floating, 30–50 cm long or more, slender. Leaves floating, 30–40 cm Χ 1–3 mm, transverse section semilunate. Racemes 5–7 cm, axis straight; staminate heads 1 or 2, smaller; pistillate heads 2 or 3, lowest one pedunculate. Male flowers: perianth segments 2–2.5 mm, attenuate toward base; filaments short; anthers oblong, ca. 1 Χ 0.5 mm. Female flowers: perianth segments linear to obtriangular, ca. 1.5 Χ 1–1.3 mm, serrate-lobate at apex; ovary elliptical; styles absent or inconspicuous; stigmas elliptical, ca. 1 mm. Fruit broadly obovoid, ca. 4 mm. 2n = 30.

Lakes, ponds, swamps; ca. 1500 m. Heilongjiang, Jilin [Japan, Russia; Europe, North America].


[1] Herbarium of Wuhan University, Department of Biology, Wuchang, Hubei, People’s Republic of China.

[2] Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, England, United Kingdom.