水玉簪科 shui yu zan ke
Wu Te-lin1, Zhang Dianxiang1; Richard M. K. Saunders2
Herbs, annual or perennial, small, mycotrophic, semi-mycotrophic, or autotrophic, often rhizomatous or tuberous. Leaves alternate, simple, entire; autotrophic species with basal rosette leaves; all with cauline leaves. Inflorescences terminal, many-flowered cymes or racemes, or flower solitary. Flowers bisexual. Perianth of 1 or 2 whorls and each whorl of 3 tepals, corolline, tubular or campanulate. Perianth tube often 3-angled or 3-winged; tepals sometimes appendaged; appendages terminal, elongated, slender. Stamens 3 or 6, if 3 then subsessile in the perianth throat, if 6 then pendent in perianth tube; connectives large, often appendiculate. Ovary inferior, 1-loculed with parietal placentation or 3-loculed with axile placentation; ovules numerous, anatropous, bitegmic; style filiform, shortly cylindrical, or conical; stigmas 3, sometimes connate. Fruit capsular, occasionally fleshy, with either persistent perianth tube and style or only persistent basal ring of perianth, dehiscence irregular or by transverse ventral slits. Seeds small, numerous; endosperm present.
About 16 genera and 148 species; tropical and subtropical regions; three genera and 13 species (two endemic) in China.
1a.... Perianth tube urceolate, circumscissile; style very short; stamens 6 ............................................................ 3. Thismia
1b.... Perianth tube salverform, cylindrical, trigonous, or campanulate; style ± as long as perianth tube; stamens 3.
2a.... Perianth persistent after anthesis; ovary 3-loculed with axile placentation ........................................... 1. Burmannia
2b.... Perianth caducous after anthesis; ovary 1-loculed with parietal placentation ................................. 2. Gymnosiphon
1. BURMANNIA Linnaeus, Sp. Pl. 1: 287. 1753.
水玉簪属 shui yu zan shu
Herbs, annual or perennial, mycotrophic, semi-mycotrophic, or autotrophic, sometimes rhizomatous. Stems simple, rarely branched. Basal leaves in a distinct rosette; cauline leaves bractlike, small. Flowers solitary or in terminal cymes. Perianth tube actinomorphic, cylindrical or trigonous, with 3 lateral wings or sometimes wings absent, without annulus at tube mouth; tepals generally 6, apically distinct; outer tepals larger than inner, unlobed; inner tepals often minute, occasionally lacking. Stamens 3, sessile or subsessile on perianth tube; connective with 02 apical and 01 basal appendages. Ovary trigonous, 3-loculed, with axile placentation; stigmas 2-labiate to funnelshaped. Fruit capsular, with persistent perianth, dehiscence generally irregular.
About 57 species; tropical and subtropical regions; 10 species (one endemic) in China.
1a.... Plants with chlorophyll, autotrophic or semi-mycotrophic; basal rosette leaves present.
2a.... Basal leaves numerous; cauline leaves 1.58 cm; inflorescences 820-flowered .................................. 1. B. disticha
2b.... Basal leaves few; cauline leaves 0.53 cm; inflorescences 16(8)-flowered.
3a.... Stems to 40 cm tall; outer tepal margins double and prominent; inner tepal margins double ............. 2. B. coelestis
3b.... Stems to 24 cm tall; outer tepal margins single or double and shallow; inner tepal margins single.
4a.... Inflorescences 13(or 4)-flowered; flowers 38 mm; stamens sessile in perianth tube ...................... 3. B. chinensis
4b.... Inflorescences (1)35(8)-flowered; flowers 914 mm; stamens with short filament, basally adnate to perianth tube ....................................................................................................................................................................... 4. B. filamentosa
1b.... Plants without chlorophyll, holomycotrophic; basal rosette leaves absent.
5a.... Stems, leaves and flowers ± purplish blue.
6a.... Perianth wings narrow or reduced to ribs; outer tepals ca. 1 mm ........................................................... 7. B. wallichii
6b.... Perianth wings conspicuous, 12 mm wide; outer tepals 1.52 mm ........................................................... 8. B. itoana
5b.... Stems, leaves and flowers whitish or yellow.
7a.... Inflorescences (1)313-flowered; inner tepals present, sometimes very small.
8a.... Perianth wings conspicuous, 12 mm wide; stamen connectives with a pendent basal spur; seeds fusiform 5. B. nepalensis
8b.... Perianth wings reduced to ribs or absent; stamen connectives without a pendent basal spur; seeds subglobose 10. B. championii
7b.... Inflorescences 14-flowered; inner tepals absent.
9a.... Stems to 20 cm; perianth tube 4.55.5 mm; perianth wings 35.5 mm wide; stamen connectives with 2 apical crests; seeds fusiform .................................................................................................................................................. 6. B. oblonga
9b.... Stems to 9 cm; perianth tube 34 mm; perianth wings 0.52.5 mm wide; stamen connectives with 1 apical crest; seeds ellipsoid ..................................................................................................................................................... 9. B. cryptopetala
1. Burmannia disticha Linnaeus, Sp. Pl. 1: 287. 1753.
水玉簪 shui yu zan
Plants with chlorophyll, autotrophic, flowering when 12.570 cm tall. Stems green, robust. Basal rosette leaves numerous, linear-lanceolate, 0.21.7 X 0.32 cm. Cauline leaves linear, 1.58 X 0.21 cm. Inflorescences 820-flowered; bracts lanceolate, 0.52.4 cm. Flowers pedicellate, bluish purple or yellow, 12.6 cm. Perianth tube 38 mm; wings conspicuous, 13.5 mm wide; outer tepals triangular, 1.54.5 mm, margin double and shallow; inner tepals lanceolate, 12 mm, margin single or double and shallow. Stamens sessile; connectives with 2 apical crests and a pendent basal spur. Ovary ellipsoid, 612 mm. Capsule obovoid. Seeds ellipsoid. Fl. AprJun, fr. JunSep. 2n = ca. 136.
Wet thickets or grassland; 4003000 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Yunnan [Cambodia, India, Indonesia, Laos, Malaysia, Myanmar, Nepal, New Guinea, Sri Lanka, Thailand, Vietnam; Australia].
2. Burmannia coelestis D. Don, Prodr. Fl. Nepal. 44. 1825.
三品一枝话 san pin yi zhi hua
Plants with chlorophyll, semi-mycotrophic, flowering when 1040 cm tall. Stems green, slender. Basal rosette leaves few, linear-lanceolate, 0.52 cm X ca. 3 mm. Cauline leaves linear, 0.53 X 0. 20.3 cm. Inflorescences (1 or)26(8)-flowered; bracts lanceolate, 36 mm. Flowers pedicellate to subsessile, bluish purple with yellow tepals, 11.5 cm. Perianth tube 4.513 mm; wings conspicuous, 1.53 mm wide; outer tepals ovate-triangular, 12 mm, margin double and prominent; inner tepals triangular, 0.51 mm, margin double. Stamens sessile; connectives with 2 apical crests and a pendent basal spur. Ovary ellipsoid to obovoid, 37 mm. Capsule obovoid. Seeds ellipsoid. Fl. JunOct, fr. OctFeb. 2n = ca. 16.
Generally in wet grassland; 0300(800) m. Guangdong, Guangxi, Hainan [Bangladesh, Cambodia, India, Indonesia, Laos, Malaysia, Myanmar, Nepal, New Guinea, Thailand, Vietnam; Australia].
3. Burmannia chinensis Gandoger, Bull. Soc. Bot. France 66: 290. 1919.
香港水玉簪 xiang gang shui yu zan
Burmannia pusilla (Wallich ex Miers) Thwaites var. hongkongensis Jonker.
Plants with chlorophyll, semi-mycotrophic, flowering when 4.520 cm tall. Stems green, slender. Basal rosette leaves few, lanceolate, 0.31.2 cm X 13 mm. Cauline leaves linear, 518 X ca. 2 mm. Inflorescences 13(or 4)-flowered; bracts lanceolate, 56 mm. Flowers pedicellate to subsessile, purple or yellow, 38 mm. Perianth tube 46.5 mm; wings conspicuous, 13 mm wide; outer tepals broadly ovate, 13 mm, margin single or double and shallow; inner tepals spatulate, 0.51.2 mm, margin single. Stamens sessile; connectives with 2 apical crests and sometimes a pendent basal spur. Ovary ellipsoid to obovoid, 2.57 mm. Capsule obovoid. Seeds ellipsoid. Fl. JunOct, fr. JulJan. 2n = ca. 32.
Generally in wet grassland; (0)3001300 m. Fujian, Guangdong, Guangxi, Hainan, Hunan, Jiangxi, Yunnan, Zhejiang [India, Laos, Thailand, Vietnam].
4. Burmannia filamentosa D. X. Zhang & R. M. K. Saunders, Nordic. J. Bot. 20: 392. 2000.
粤东水玉簪 yue dong shui yu zan
Plants with chlorophyll, semi-mycotrophic, flowering when 824 cm tall. Stems green, slender. Basal rosette leaves few, triangular-lanceolate, 0.51.4 cm X 23 mm. Cauline leaves linear, 515 X 1.53 mm. Inflorescences (1)35(8)-flowered; bracts lanceolate, ca. 5 mm. Flowers subsessile, bluish purple, 0.91.4 cm. Perianth tube 67.5 mm; wings conspicuous, 23 mm wide; outer tepals narrowly triangular, 1.53 mm, margin double and shallow; inner tepals lanceolate, 12 mm, margin single. Stamens with short filament, basally adnate to perianth tube; connectives with 2 apical crests but lacking a pendent basal spur. Ovary ellipsoid, 37 mm. Capsule obovoid. Seeds ellipsoid. Fl. and fr. OctNov.
* Wet, open areas; 300800 m. E Guangdong.
5. Burmannia nepalensis (Miers) J. D. Hooker, Fl. Brit. India 5: 666. 1888.
宽翅水玉簪 kuan chi shui yu zan
Gonyanthes nepalensis Miers, Trans. Linn. Soc. London 18: 537. 1841; Burmannia fadouensis H. Li; B. liukiuensis Hayata.
Plants without chlorophyll, holomycotrophic, flowering when 413 cm tall. Stems whitish, slender. Basal rosette leaves absent. Cauline leaves triangular, 23.5 X ca. 1 mm. Inflorescence (1)39-flowered; bracts triangular-lanceolate, 23 mm. Flowers pedicellate, whitish or yellow, 47 mm. Perianth tube 1.52.5 mm; wings conspicuous, 12 mm wide; outer tepals oblong-ovate, 0.51.2 mm, margin single; inner tepals orbiculate, very small, margin single. Stamens sessile; connectives with 2 apical crests and a pendent basal spur. Ovary subglobose, 1.53 mm. Capsule subglobose. Seeds fusiform. Fl. and fr. JulDec.
Wet areas in forests; 4001600 m. Fujian, Guangdong, Guangxi, Hunan, Taiwan, Yunnan [India, Indonesia, Japan, Nepal, Philippines, Thailand].
6. Burmannia oblonga Ridley, J. Straits Branch Roy. Asiat. Soc. 41: 33. 1904.
裂萼水玉簪 lie e shui yu zan
Burmannia bifida Gagnepain.
Plants without chlorophyll, holomycotrophic, flowering when 720 cm tall. Stems whitish, slender. Basal rosette leaves absent. Cauline leaves triangular, 1.53 X 12 mm. Inflorescences 14-flowered; bracts triangular-lanceolate, 2.53 mm. Flowers pedicellate to subsessile, whitish, 0.81.4 cm. Perianth tube 4.55.5 mm; wings conspicuous, 35.5 mm wide; outer tepals broadly oblong, ca. 2 mm, margin single; inner tepals absent. Stamens sessile; connectives with 2 apical crests and a pendent basal spur. Ovary globose to rhomboid, ca. 4 mm. Capsule subglobose. Seeds fusiform. Fl. and fr. JulDec.
Wet, evergreen forests; 8001100 m. Hainan [Cambodia, Indonesia, Malaysia, Thailand, Vietnam].
7. Burmannia wallichii (Miers) J. D. Hooker, Fl. Brit. India 5: 666. 1888.
亭立 ting li
Gonyanthes wallichii Miers, Trans. Linn. Soc. London 18: 537. 1841.
Plants without chlorophyll, holomycotrophic, flowering when 411.5 cm tall. Stems purple, slender. Basal rosette leaves absent. Cauline leaves triangular-subulate, 1.53.5 X 11.5 mm. Inflorescences 13-flowered; bracts triangular-lanceolate, 23.5 mm. Flowers sessile, bluish purple, 510 mm. Perianth tube 2.55.5 mm; wings narrow or reduced to ribs, 0.20.8 mm wide; outer tepals ovate-triangular, ca. 1 mm, margin single; inner tepals orbiculate, very small, margin single. Stamens sessile; connectives lacking apical crests, sometimes with a pendent basal spur. Ovary ellipsoid to subglobose, 1.53 mm. Capsule subglobose. Seeds ellipsoid. Fl. and fr. JulNov. 2n = 32.
Wet areas in dense forests; 0700 m. Guangdong, Hainan, Yunnan [Myanmar, Thailand, Vietnam].
8. Burmannia itoana Makino, Bot. Mag. (Tokyo) 27: 1. 1913.
纤草 xian cao
Burmannia pingbienensis H. Li; B. takeoi Hayata.
Plants without chlorophyll, holomycotrophic, flowering when 515 cm tall. Stems light blue, slender. Basal rosette leaves absent. Cauline leaves ovate-lanceolate, 2.53.5 X 1.52.5 mm. Inflorescence 13-flowered; bracts ovate-lanceolate, ca. 5 mm. Flowers pedicellate, bluish purple, 811 mm. Perianth-tube 3.55.5 mm; wings conspicuous, 12 mm wide; outer tepals ovate-triangular, 1.52 mm, margin single; inner tepals orbiculate, very small, margin single. Stamens sessile; connectives with an apical crest and a pendent basal spur. Ovary subglobose, 23.5 mm. Capsule subglobose. Seeds ellipsoid. Fl. and fr. JunDec.
Wet evergreen forests; 3001200 m. Guangdong, Guangxi, Hainan, Taiwan, Yunnan [Japan].
9. Burmannia cryptopetala Makino, Bot. Mag. (Tokyo) 27: 3. 1913.
透明水玉簪 tou ming shui yu zan
Burmannia cryptopetala Makino var. daxikangensis Z. Wei & Y. B. Chang.
Plants without chlorophyll, holomycotrophic, flowering when 59 cm tall. Stems whitish, slender. Basal rosette leaves absent. Cauline leaves ovate-lanceolate, 33.5 X 1.52.5 mm. Inflorescences 13-flowered; bracts ovate-lanceolate, ca. 3 mm. Flowers pedicellate, whitish with yellow tepals, 11.2 cm. Perianth tube 34 mm; wings conspicuous, 0.52.5 mm wide; outer tepals triangular, 1.52 mm, margin single; inner tepals absent. Stamens sessile; connectives with an apical crest and a pendent basal spur. Ovary subglobose, 3.55 mm. Capsule subglobose. Seeds ellipsoid. Fl. and fr. JunNov.
Wet areas in dense forests; 200800 m. Guangdong, Hainan, Zhejiang [Japan].
10. Burmannia championii Thwaites, Enum. Pl. Zeyl. 325. 1864.
头花水玉簪 tou hua shui yu zan
Burmannia dazielii Rendle; B. hunanensis K. M. Liu & C. L. Long.
Plants without chlorophyll, holomycotrophic, flowering when 322 cm tall. Stems whitish, slender. Basal rosette leaves absent. Cauline leaves ovate-triangular, 1.56 X 13 mm. Inflorescences 313-flowered; bracts ovate-triangular, 1.56 mm. Flowers pedicellate or sessile, whitish with yellow tepals, 511 mm. Perianth tube 35 mm; wings reduced to ribs or absent; outer tepals narrowly triangular, 1.53 mm, margin single; inner tepals spatulate, 0.81.4 mm, margin single. Stamens sessile or with a short filament, basally adnate to perianth; connectives sometimes with 2 small apical crests but lacking a pendent basal spur. Ovary ellipsoid to obovoid, 2.53.5 mm. Capsule subglobose. Seeds subglobose. Fl. and fr. MayAug.
Wet areas in forests; 01700 m. Fujian, Guangdong, Guangxi, Hunan, Taiwan [India, Indonesia, Japan, Malaysia, New Guinea, Sri Lanka, Thailand; Pacific Islands].
2. GYMNOSIPHON Blume, Enum. Pl. Javae 1: 29. 1827.
腐草属 fu cao shu
Herbs, annual, without chlorophyll, mycotrophic; rhizomatous. Stems simple or branched. Leaves cauline, bractlike, small. Inflorescences terminal, racemes or flower solitary. Perianth tube actinomorphic, salverform, lateral wings absent, without annulus at mouth; tepals 6, apically distinct; outer tepals larger than inner, slightly 3-lobed; inner tepals very small. Stamens 3, sessile on perianth tube; connectives generally unappendaged. Ovary ovoid to globose, 1-loculed, with 3 parietal placentas; placentas each with a large globose gland at both sides of the apex; stigmas 3, often appendiculate. Fruit capsular, perianth caducous but perianth base persistent; dehiscence generally irregular or by longitudinal slits.
About 26 species; Africa, tropical America, Asia, New Guinea; one species in China.
1. Gymnosiphon aphyllus Blume, Enum. Pl. Javae 1: 29. 1827
腐草 fu cao
Burmannia nana Fukuyama & T. Suzuki; Gymnosiphon nanus (Fukuyama & T. Suzuki) Tuyama.
Plants holomycotrophic, flowering when 78 cm tall. Stems white to pale yellow. Leaves narrowly ovate-triangular, 1.52 mm. Inflorescence branched or not, 58-flowered; bracts ovate-triangular, ca. 2 mm. Flowers pedicellate, white; perianth tube cylindrical, ca. 3 mm; outer tepals ovate, ca. 1 mm; inner tepals ca. 0.2 mm. Stamens inserted immediately below inner tepals. Stigmas curved, funnel-shaped, unappendiculate. Fruit ca. 3 mm, dehiscence by irregular perforation of the wall between costae. Seeds ovoid.
Tropical rain forests; low elevations. Taiwan [Indonesia, Malaysia, New Guinea, Thailand].
3. THISMIA Griffith, Proc. Linn. Soc. Lond. 1: 221. 1844.
水玉杯 shui yu bei
Plants annual, without chlorophyll, mycotrophic; tuberous. Stems simple or branched. Leaves cauline, bractlike, small. Flowers solitary, terminal. Perianth tube actinomorphic to zygomorphic, cylindrical, urceolate, or sometimes campanulate, with prominent annulus at mouth, lateral wings absent; tepals 6, apically distinct; outer tepals as long or shorter than inner tepals; inner tepals sometimes apically connivent or connate into an erect mitre. Stamens 6, generally laterally connate, pendent from annulus, occasionally distinct; connectives with 02 apical and 02 basal appendages. Ovary obconical or obovoid, 1-loculed, with 3 parietal placentas; stigmas 3, simple or bilabiate. Fruit fleshy with persistent perianth base; dehiscence by abscission of apex.
About 43 species; pantropical; two species (one endemic) in China.
1a.... Flowers entirely white; inner tepal appendage 2.83.3 cm; outer tepal appendage 58 mm ......... 1. T. taiwanensis
1b.... Flowers white with yellow annulus and orangish red tepal appendages; inner tepal appendage ca. 1.7 cm; outer tepal appendage absent ...................................................................................................................................... 2. T. tentaculata
1. Thismia taiwanensis S. Z. Yang et al., Syst. Bot. 27: 485. 2002.
台湾水玉杯 tai wan shui yu bei
Plants holomycotrophic. Stems ± absent at anthesis, 1.34 cm and carnose in fruit. Leaves scalelike, 3.35.5 X 0.41.0 mm. Flowers solitary, subsessile, with 2 or 3 subtending bracts. Perianth tube white, translucent, actinomorphic, urceolate, ca. 9 X 5 mm, circumscissile. Outer tepals ovate, 2.53.5 mm excluding appendage, radiating outwards; appendage white, erect, subulate, 58 mm. Inner tepals spatulate, ca. 5 mm excluding appendage, apically imbricate and forming a loose mitre over perianth chamber; appendage white, erect, subulate, 2.83.3 cm. Stamens apparently distinct. Fruit capsule white, translucent, cup-shaped, 3.33.7 X ca. 4 mm, apically dehiscent; fruiting peduncle 58 cm. Seeds ellipsoid. Fl. and fr. Sep.
* Fagaceae forests; 20002100 m. Taiwan (Gaoxiong).
2. Thismia tentaculata K. Larsen & Averyanov, Rheedea 17: 16. 2007.
三丝水玉杯 san si shui yu bei
Plants holomycotrophic. Stems white, unbranched, erect, 2.53 cm. Leaves scalelike, 1.55.7 X 0.81.1 mm. Flowers 14, subsessile, with subtending bracts. Perianth tube white, translucent, actinomorphic, urceolate, ca. 7.3 X 4.8 mm, circumscissile. Outer tepals broadly triangular, ca. 1.4 mm, lacking appendage. Inner tepals narrowly triangular, ca. 2.3 mm excluding appendage; appendage orangish red, spreading, filiform, ca. 1.7 cm. Stamens apically connate. Fruit capsule white, translucent, cup-shaped. Seeds ellipsoid. Fl. and fr. JunAug.
Evergreen montane forests; 600800 m. Hong Kong [Vietnam].