OPHIOGLOSSACEAE [Draft]

瓶尔小草科 ping er xiao cao ke

Liu Quanru (刘全儒); N. Sahashi

Plants terrestrial, rarely epiphytic, generally small, erect, rarely pendent. Rhizomes short, erect, bearing a cluster of stout fibrous roots and one to several fronds. Fronds erect or pendent, dimorphic, consisting of sterile (trophophylls) and fertile (sporophylls) portions, both arising from common stalk. Sterile blades 1 or 2, rarely more, simple or once or twice forked at apex, lanceolate, ribbon-shaped, or ovate, with entire or wavy margin; veins reticulate, with indistinct midrib. Fertile frond portion branching from sterile blade. Sporangia embedded in a linear spike, arranged in two rows along margin, dehiscing by transverse slits; spores tetrahedral, trilete, globose or rounded triangular, surface irregularly reticulate or foveo-reticulate or foveolate with fine granules. [??x = ]. 2n = 240–1440.

Four genera and ca. 30 species: widespread; nine species in China (one endemic).

1a.     Plants terrestrial; sterile blades small, ovate or lanceolate, erect; rhizomes glabrous; fertile frond portion arising from base of sterile blade ...........................................................................  1. Ophioglossum

1b.     Plants epiphytic; sterile blades large, ribbon-shaped, pendulous; rhizome with long hairs; fertile frond portion arising medially from or toward base of sterile blade ..................................................  2. Ophioderma

 

1. OPHIOGLOSSUM Linnaeus, Sp. Pl. 2: 1062. 1753.

瓶耳小草属  ping er xiao cao shu

Ferns terrestrial, small, erect. Fronds dimorphic: sterile blades 1 or 2, rarely more, with stalks, usually simple, lanceolate or oval, with entire or wavy margin; veins reticulate, with indistinct midrib; fertile frond portion arising from base of sterile blade, with long stalk.

About 25 species; mainly in the Northern Hemisphere; eight species in China (one endemic).

[??Ophioglossum japonicum Thunberg]

[Fl. Fujian.: ??Ophioglossum flexuosum]

[Fl. Fujian, Jiangxi: ??Ophioglossum scandens]

[Fl. Fujian: ??Ophioglossum zeylanicum]

1a.     Plants usually 2–10 cm tall; sterile blade usually less than 3 cm.

2a.    Fronds consisting of sterile and fertile portions, sterile blade elliptic or elliptic-ovate; spike bearing 10–18 pairs of sporangia ................................................................................................. 7. O. nudicaule

2b.    Fronds usually consisting of fertile portion only, or rarely with both sterile and fertile portions; sterile blade elliptic or ovate-rhombic; spike bearing 3–12 pairs of sporangia ................  8. O. kawamurae

1b.     Plants more than 10 cm tall, sometimes up to 35 cm; sterile blade more than 3 cm.

3a.    Sterile blade cordate at base; margin ± wavy; areolae of veins evident.

4a.     Spores coarsely reticulate .........................................................  4. O. austro-asiaticum

4b.     Spores subreticulate ..........................................................................  5. O. reticulatum

3b.    Sterile blade not cordate at base; margin entire.

5a.     Sterile blade narrow, lanceolate or oblanceolate, narrowly cuneate at base  6. O. thermale

5b.     Sterile blade broad, oblong to ovate, broadly cuneate, truncate, or rounded at base.

6a.     Sterile blade oblong, rounded at apex ...............................................  3. O. oblongum

6b.     Sterile blade elliptic or narrowly to broadly ovate, rounded or acute at apex.

7a.    Sterile blade elliptic or narrowly ovate, 6–10 cm or longer ..............  1. O. vulgatum

7b.    Sterile blade ovate to broadly ovate, 3–6 cm ................................  2. O. petiolatum

 

1. Ophioglossum vulgatum Linnaeus, Sp. Pl. 2: 1062. 1753.

瓶尔小草  ping er xiao cao

[??Ophioglossum nipponicum Miyabe & Kudo]

Plants 10–30 cm tall. Rhizomes erect, bearing a cluster of fleshy roots. Roots extended horizontally and stolonlike. Fronds usually single; common stalk pale at lower part, 6–9 cm, slightly robust, deeply buried underground. Sterile blade without a stalk, elliptic or narrowly ovate, rarely ovate, 6–10 Χ 1.5–4 cm, slightly fleshy to herbaceous, base cuneate, truncate, or rounded, sometimes abruptly narrowed, margin entire, apex obtuse or abruptly acute; veins distinctly reticulate. Fertile frond portion arising from base of  sterile blade, 9–18 cm or longer; spike 2.5–3.5 cm, ca. 0.2 mm in diam., apex acute, exceeding sterile blade; spore surface obviously and coarsely reticulate. 2n = 240–960.

Shaded forests, wet meadows; up to 3000 m. Anhui, Fujian, Guangdong, Guizhou, [??Henan], Hubei, Hunan, Jiangxi, S Shaanxi, Sichuan, Xizang, Yunnan, Zhejiang [India, Japan, Korea, Sri Lanka; Australia, North America, Europe].

2. Ophioglossum petiolatum Hooker, Exotic Fl. 1: t. 56. 1823.

柄叶瓶尔小草  bing ye ping er xiao cao

Plants 15–25 cm tall. Rhizomes erect, bearing a cluster of thick fleshy roots, which extend horizontally like stolons, producing a new plant from apical bud. Fronds simple; common stalk 9–15 cm. Sterile blade sessile, ovate to broadly ovate, 3–6 Χ 2–3 cm, herbaceous, base ± elongated and rounded, apex acute or obtuse; veins fairly distinct, reticulate. Fertile frond portion arising from base of sterile blade, 6–9 cm, linear spike 2.5–3 cm. Spore surface obviously subreticulate with fine granules. 2n = 240–960.

Open shrubby hillsides; 200–3300 m. Fujian, Guizhou, [??Hainan: specimen of O. pedunculosum at A], Hubei, Sichuan, Taiwan, Yunnan [W India, Indonesia, Japan, Nepal, Philippines, Sri Lanka, Thailand; Australia, North America, New Zealand].

Specimens described as Ophioglossum pedunculosum Desvaux in Flora Reipublicae Popularis Sinicae (2: 9–10. 1959) and some provincial floras (e.g., floras of Anhui, Fujian, Guangdong, Guizhou, Jiangxi, Yunnan) should be included in O. petiolatum. According to the studies by Clausen (1938 [??full citation]), O. pedunculosum was described as having a blade with a prominent pale median band, but this character is very different in Chinese specimens. Whether O. pedunculosum is really distributed in China needs further study.

3. Ophioglossum oblongum H. G. Zhou & H. Li, Guihaia 1: 40. 1991.

矩圆叶瓶尔小草  ju yuan ye ping er xiao cao

Plants 20–25 cm tall. Rhizomes erect, bearing few fleshy roots. Fronds caespitose, ca. 23 cm; common stalk 6–8 cm, greenish or pale at lower part from being buried underground. Sterile blade simple, sessile, light green when dried, oblong, 2–8 Χ ca. 1.5 cm, herbaceous, base broadly cuneate, shortly decurrent, margin entire, apex rounded; reticulate veins indistinct. Fertile frond portion 6–14 cm, slender, arising from base of sterile blade; spike linear, 1.5–4 cm, bearing 17–34 pairs of sporangia.

* Guangxi (Fengshan).

4. Ophioglossum austro-asiaticum Nishida, Bull. Nat. Sci. Mus. 4: 329. 1959 [??J. Jap. Bot. 34: 46, t. 2(A). 1959].

高山瓶尔小草  gao shan ping er xiao cao

Plants 10–20 cm tall. Rhizomes erect, cylindrical. Common stalk 6.5–7.5 cm. Sterile blade sessile, nearly orbicular or broadly ovate, ca. 3.5 Χ 3 cm, base cordate, apex rounded; areolae with included veinlets and smaller areolae. Fertile segment arising from near base of sterile blade. Spore surface coarsely reticulate.

Taiwan [Indonesia (Borneo)].

Ophioglossum austro-asiaticum resembles O. reticulatum, but differs from the latter in having coarsely reticulate exospores only. This species  needs further study.

5. Ophioglossum reticulatum Linnaeus, Sp. Pl. 2: 1063. 1753.

心叶瓶尔小草  xin ye ping er xiao cao

[??Ophioglossum cordifolium Roxburgh; O. pedunculosum Dunn & Tutcher (1912), not Desvaux (1811)]

Plants 10–30 cm tall. Rhizomes erect, slender, bearing a few thick fleshy roots. Common stalk 4–8 cm, light green, gradually pale toward base. Sterile blade with a short stalk or almost sessile, ovate or orbicular-ovate, 3–4 Χ 2.6–3.5 cm, herbaceous, base deeply cordate, margin ± wavy, apex rounded or subobtuse; veins distinct, reticulate. Fertile frond portion arising from base of sterile blade, slender, 10–15 cm; spike 3–3.5 cm, slender. Spore surface regularly or irregularly subreticulate. 2n = 480–1440.

Shaded forests; 1100–4000 m. Fujian, Gansu, Guizhou, Henan, [??Hubei: specimen at A], Jiangxi, Shaanxi, Sichuan, [??Taiwan: Taiwania 30: 53. 1985], Xizang, Yunnan [Korea; South America].

6. Ophioglossum thermale Komarov, Repert. Spec. Nov. Regni Veg. 13: 85. 1914.

狭叶瓶尔小草  xia ye ping er xiao cao

[??Ophioglossum angustatum Maxon; O. japonicum Prantl (1883), not Thunberg (1784); O. nipponicum Nakai (1926), not Miyabe & Kudo (1916); O. nudicaule H. Christ (1896), not Linnaeus f. (1781); O. savatieri Nakai; O. vulgatum var. thermale C. Christensen]

Plants 10–20 cm tall. Rhizomes erect, slender, bearing a cluster of unbranched fleshy roots; roots stolonlike, producing new plants at apices. Fronds simple or 2 or 3 together. Common stalk green or pale when lower part buried underground, 3–6 cm, slender. Sterile blade sessile, light green, simple, oblanceolate or oblong-oblanceolate, 2–5 cm Χ 3–10 mm, herbaceous, base narrowly cuneate, margin entire, apex slightly acute or obtuse; veins reticulate, indistinct, but visible under light. Fertile frond portion arising from base of sterile blade; stalk 5–7 cm, overtopping sterile blade; spike narrowly linear, 2–3 cm, acute at apices, with 15–28 pairs of sporangia. Spores pale, surface subreticulate or nearly smooth. 2n = 240–720.

Grassy hillsides, near thermal springs; 200–1800 m. [??Anhui – specimen of O. angustatum at A], Hebei, Heilongjiang, Henan, Jiangsu, Jiangxi, Jilin, Liaoning, Shaanxi, [??Shandong], Sichuan, Yunnan [Japan, Korea, Russia (Kamchatka)].

7. Ophioglossum nudicaule Linnaeus f., Suppl. Pl. 443. 1781.

裸茎瓶尔小草  luo jing ping er xiao cao

Ophioglossum parvifolium Greville & Hooker.

Plants 3–8 cm tall. Rhizomes erect, very thick, with numerous fibrous roots, usually bearing 2 or 3 fronds. Common stalk grayish green on most parts from being buried underground, 1.5–3 cm. Sterile blade elliptic or elliptic-ovate, 1–3 Χ ca. 1 cm, subfleshy, base shortly cuneate, margin entire, apex acute or rounded; veins indistinct. Fertile frond portion 4–5 cm, arising from base of sterile blade; spike short and thick, 1–2 cm, surpassing sterile blade 1–2 times, apex acute; sporangia 10–18 pairs. Spore surface coarsely reticulate. 2n = 240–1000.

Slopes, meadows; 1800–4300 m. SW Sichuan, Xizang, C and NW Yunnan [N India, Japan, Nepal].

8. Ophioglossum kawamurae Tagawa, Acta Phytotax. Geobot. 8: 134. 1939.

永仁瓶尔小草  yong ren ping er xiao cao

Ophioglossum yongrenense Ching ex Z. R. He & W. M. Chu.

Plants 2–9 cm tall. Rhizomes erect, cylindrical, with fleshy roots, 1 or 2 new fronds arising every year. Common stalk 2–10 mm, ca. 0.5 mm in diam. Fronds usually consisting only of fertile blades, rarely with both sterile and fertile blade portions. Sterile blade sessile, adpressed to ground, elliptic or ovate-rhomboid, 0.5–1.5 Χ 0.2–0.7 cm, apex abruptly acute. Spike simple, rarely forked, 0.3–1.5 cm, ca. 0.1 mm [??] in diam.; stalk 0.5–6 cm, bearing 3–12 pairs of sporangia. Spores tetrahedral, globose, surface foveo-reticulate. 2n = 240.

Dry slopes, grasslands; ca. 2000 m. Yunnan (Yongren) [Japan].

2. OPHIODERMA (Blume) Endlicher, Gen. Pl. 66. 1836.

带状瓶耳小草属  dai zhuang poing er xiao cao shu

Ophioglossum sect. Ophioderma Blume, Enum. Pl. Jav. 2: 259. 1828.

Plants large, epiphytic. Rhizomes short, with fleshy roots. Sterile blade ribbon-shaped, long, pendulous, base narrowed, apex usually forked; veins reticulate. Fertile frond portion spikelike, arising from middle part of sterile blade, simple, sometimes branched. Sporangia immersed in central axis of fertile frond segment.

Three species; mainly in tropical rainforests; one species in China.

1. Ophioderma pendula (Linnaeus) C. Presl, Suppl. Tent. Pterid. 56. 1845.

带状瓶尔小草  dai zhuang ping er xiao cao

Ophioglossum pendulum Linnaeus, Sp. Pl. 2: 1518. 1763; [??Ophioderma pendula f. ramosa Nakai].

Fronds 1–3, pendulous and ribbon-like, usually lanceolate, up to 30–50 or more Χ 1–3 cm, sessile; veins reticulate, without distinct midrib, forming diagonal and hexagonal areoles; veinlets ± visible. Fertile frond portion arising from near base or middle part of sterile blade and never longer than sterile blade; spike 5–25 cm; sporangia numerous, 40–200 on each side. Spores achromatous [??without color or white] or light yellow, quadrangular, surface foveolate (small pits scattered throughout surface). 2n = 740–960.

On trunks of trees in tropical rainforests. Hainan, Taiwan, Yunnan [India, Indonesia, Japan, Korea, Malaysia, Philippines, Sri Lanka; W Africa, Australia, Polynesia (Hawaiian islands)].