SPARGANIACEAE [draft]

黑三棱科  hei san leng ke

Sun Kun (孙坤)[1]; David A. Simpson[2]

Perennial aquatic or marsh herbs with creeping rhizomes. Stems simple or branched, robust or slender. Leaves alternate, distichous, emergent or floating, sheathing at base. Flowers unisexual, monoecious, 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 borne 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-celled or rarely 2-celled, narrow at base; styles simple or forked, with a unilateral stigma; ovule 1, pendulous. Fruit nutlike, indehiscent. Seeds with thin testa.

One genus and about 19 species: mainly in temperate and cool regions of the world, best represented in N 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.

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 obovate .....................  11. S. hyperboreum

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

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

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

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

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

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

6b.       Panicles with 1(–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 to one pistillate heads 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 a inconspicuous stalk  4. S. fallax

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

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

10b.     Staminate heads 1–3, not separated 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 erect, robust, 70–120 cm tall or more, emersed. Leaves (20–)40–90 Χ 0.7–1.6 cm, upper flattened, lower keeled or trigonous on abaxial side. Panicles 20–60 cm long, with 3–7 lateral branches, each with 7–11 staminate and 1–2 pistillate heads; main axis consisting of 3–5 or more staminate heads. Perianth segments of male flowers spathulate, lobate at apex; filaments ca. 3 mm; anthers obconical, 1–1.2 Χ 0.5 mm; perianth segments of female flowers 5–7 Χ 1–1.5 mm; ovary sessile; styles forked or simple, ca. 1.5 mm; stigmas 3–4 mm. Fruit obconical, 6–9 mm long, ribbed. Fl. and fr. May–Oct.

In lakes, streams, swamps and ponds, up to ca. 3600 m. Gansu, Hebei, Heilongjiang, Hubei, Jiangsu, Jiangxi, Jilin, Liaoning, Nei Mongol, Shaanxi, Shanxi, Xinjiang, Xizang, Yunnan [Afghanistan, C Asia, Japan, Korea, Russia].

1a. Diagnosis ................................................................................................  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  1a. 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.

Diagnosis.

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 S. 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 long, or for the stems arcuate at the upper part, fruits slightly small. They represent local variant individuals rather than distinct taxa.

1b. Sparganium stoloniferum subsp. choui (D. Yu) 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 long, each branch with only one staminate heads; 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, slender, 20–36 cm tall. Leaves 25–35 cm Χ 2–3 mm, upper flattened, lower keeled or trigonous on abaxial side. Panicles 7–15 cm long, main axis consisting of 5–7 staminate heads and 2–3 pistillate heads, usually with 1 lateral branch at base; lateral branch 5–8 cm, with 2–3 staminate and 1–2 pistillate heads. Perianth segments of male flowers spathulate, ca. 2 mm, lobate at apex; filaments ca. 2 mm; anthers oblong, ca. 1 Χ 0.3 mm; perianth segments of female flowers spathulate, ca. 2 mm, lobate at apex; ovary sessile; styles ca. 0.5 mm; stigmas unilateral, ca. 1.5 mm. Fruit obovate, 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 erect, 33–57 cm tall, emersed. Leaves erect, 50–70 cm Χ 4–5 mm, upper flattened, lower keeled or trigonous on abaxial side. Panicles 7–16 cm long, with 1 lateral branch at base, main axis and lateral branch usually zigzag; main axis with 5–8 staminate heads and 2–3 pistillate heads; lateral branch with 3(–5) staminate and 1 pistillate heads. Perianth segments of male flowers ca. 1.5 Χ 0.2 mm; filaments 3–4 mm; anthers oblong, ca. 1 Χ 0.2 mm; perianth segments of female flowers 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 erect, 40–55(–80) cm tall, emersed. Leaves 45–65 cm Χ 4–10 mm, upper flattened, lower keeled or trigonous on abaxial side. Racemes 15–17 cm long, lower curved; staminate heads 4–7, distinctly separated from pistillate ones; pistillate heads 3–4(–7), relatively widely separated, sessile or the lowest pedunculate. Perianth segments of male flowers linear, 2–2.5 mm long, serrate-lobate at aex; filaments 3–4.5 mm; anthers oblong, ca. 1.5 Χ 0.4 mm; perianth segments of female flowers broadly spathulate, ca. 3 Χ 1–1.5 mm, serrate-lobate at aex; ovary attenuate toward base; styles short; stigmas 1.5–2 mm. Fruit broadly fusiformis, 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 erect, slender, 30–70 cm tall, emersed. Leaves 40–80 cm long, aerial or floating, lower keeled on abaxial side. Racemes 10–20 cm long; staminate heads 4–8, distinctly separated from pistillate ones; pistillate heads 3–4, relatively widely separated, sessile or the lowest 1–2 pedunculate. Perianth segments of male flowers linear or spathulate, 2–2.5 mm long, lobate at apex; filaments ca. 4 mm; anthers oblong, ca. 1.6 Χ 0.4 mm; perianth segments of female flowers spathulate, ca. 3.5 mm, lobate at apex; ovary with short stalk; styles ca. 1 mm; stigmas 1.5–1.8 mm. Fruit fusiformis. 2n = 30. Fl. and fr. Jun–Oct.

Margins of lakes, swamps, and streams. Gansu, Heilongjiang, Jilin, Liaoning, Nei Mongol, Xinjiang [Japan, Russia; and elsewhere too, surely??].

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 erect, 20–50 cm tall, emersed. Leaves 30–56 cm long, lower keeled or trigonous on abaxial side. Racemes 6–15 cm long; staminate heads 1–3, not separated from pistillate ones; pistillate heads 3–4(–8), relatively near to each other, lowest one pedunculate. Perianth segments of male flowers ca. 1.5 mm long, serrate-lobate at apex; filaments 3–3.5 mm; anthers oblong, ca. 1 mm; perianth segments of female flowers 2–2.5 mm, lobate at apex; ovary fusiformis, with short stalk; styles short; stigmas ca. 0.5 mm. Fruit broadly fusiformis, 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, Russia; Europe].

A specimen with 7 or 8 pistillate heads and larger flowers from Heilongjiang, which was described as Sarganium manshuricum by D. Yu (Bull. Bot. Res., Harbin 12: 255. 1992) mainly according to some quantitative characters, perhaps represents a local variant suite of specimen 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 erect, 15–45 cm tall, emersed. Leaves 25–55 cm Χ 3–5 mm lower keeled on abaxial side. Spikes with many densely arranged heads, 6–11 cm long; staminate heads (6–)8–12, deciduous; pistillate heads 1–3, sessile or rarely lowest one sterile, pedunculate. Perianth segments of female flowers broadly 3–5 Χ ca. 1.5 mm, lobate to parted at apex; ovary ellipitical, with stalk 1.5–2 mm long; styles ca. 1 mm; stigmas unilateral, 1–1.5 mm long. 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 stout, 1.3–1.5 m long, floating. Leaves floating, ca. 1.5 m Χ ca. 1 cm, flattened, spongy, lower slightly convex on abaxial side. Racemes 20–35 cm long, lower curved?; staminate heads 8—10; pistillate heads 3–4(–7), sessile or lowest 1–2 pedunculate. Perianth segments of male flowers spathulate, 3–3.5 mm long; filaments 3–4 mm; anthers linear, ca. 2 mm; perianth segments of female flowers white, spathulate, 4–5 mm, lobate at apex; ovary fusiformis, 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 slender, 30–50 cm long, floating. Leaves floating, 25–40 cm Χ ca. 4 mm, flattened. Racemes 6–16 cm long; staminate heads 2–4; pistillate heads 2–4, relatively widely seperated, sessile or lowest 1–2 pedunculate. Perianth segments of male flowers spathulate to obtriangular; filaments ca. 5 mm; anthers elliptical, 1–1.2 Χ ca. 0.5 mm; perianth segments of female flowers linear to spathulate, serrate-lobate to parted; ovary fusiformis, attenuate toward base and forming a short stalk; styles ca. 2 mm; stigmas long-elliptical. Fruit fusiformis to elliptical, ca. 4 mm. 2n = 30. Fl. and fr. Jul–Sep.

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

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

The 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 respecimen, respectively, their taxonomic position 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 slender, 30–50 cm long or more, floating. Leaves floating, 30–40 cm Χ 1–3 mm, transverse section semilunate. Racemes 5–7 cm long, axis straight; staminate heads 1–2, smaller; pistillate heads 2–3, lowest one pedunculate. Perianth segments of male flowers 2–2.5 mm long, attenuate toward base; filaments short; anthers oblong, ca. 1 Χ 0.5 mm; perianth segments of female flowers 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 obovate, 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.