CYSTOPTERIDACEAE [Draft: 01 May 2012]

冷蕨科  leng jue ke

Wang Zhongren (王中仁); Masahiro Kato[1], Chris Haufler, Kathleen Pryer[2]

Plants small to medium-sized, summer-green; rhizomes slender, creeping or ascending; costae articulate to rachis (in Gymnocarpium) or not so; lamina pinnate to 3(or 4)-pinnate-pinnatifid; veins free; sori orbicular or elongate, abaxial on veins, indusiate or exindusiate; indusia ovate-lanceolate, ovate, or orbicular, attached proximally to receptacle. x = 40, 42.

Four genera and more than 30 species: four genera and 20 species (eight endemic) in China.

Chu Wei-ming, Wang Zhong-ren, Hsieh Yin-tang & He Zhao-rong. 1999. Athyriaceae. In: Chu Wei-ming, ed., Fl. Reipubl. Popularis Sin. 3(2).

1a.       Sori exindusiate .........................................................................................  1. Gymnocarpium

1b.       Sori indusiate.

2a.       Multicellular articulate hairs present on stipe and lamina; indusia small, often hidden under sporangia      3. Acystopteris

2b.       Multicellular articulate hairs absent from stipe and lamina; indusia visible.

3a.       Lamina deltoid to lanceolate, base slightly narrowed or broadest part of lamina; spore wall echinate    4. Cystopteris

3b.       Lamina oblong-lanceolate, base gradually narrowed, spore wall with conical spines  2. Cystoathyrium

1. GYMNOCARPIUM Newman, Phytologist 4: 371. 1851.

羽节蕨属  yu jie jue shu

Wang Zhongren (王中仁); Kathleen Pryer

Carpogymnia (H. P. Fuchs ex Janchen) Α. Lφve & D. Lφve; Currania Copeland; Thelypteris sect. Carpogymnia H. P. Fuchs ex Janchen.

Plants terrestrial, summer-green, small to medium-sized. Rhizomes long creeping, blackish brown, glabrate, clothed with brown, thin, broadly lanceolate or ovate-lanceolate scales at apices and stipe bases. Fronds distant; stipe thin, much longer than lamina, dark brown at base, upper part stramineous, U-shaped grooved adaxially; lamina simple-pinnatipartite to 3-pinnate-pinnatifid, deltoid-ovate to pentagonal-oval, bases articulate to stipe apex, acuminate at apex; pinnae stalked or sessile, articulate to rachis, basal pair not shortened. Veins free, pinnate in ultimate segments, lateral veins simple or occasionally forked, terminating at margin. Lamina herbaceous or thinly herbaceous, stipe apex, rachis, costae, and lamina ± with hyaline or pale yellow glands on surfaces, or absent. Sori oblong or orbicular, exindusiate, abaxial on veins, uniseriate along each side of costule or midrib. Spores bean-shaped, perispore surface rugate, folds lobed, foveolate or sometimes reticulate. x = 40.

Two sections, ten species and several hybrids: distributed in the temperate zone of the N Hemisphere (Asia, North America, and Europe) and the subtropical mountains of Asia, occurring in forest understories; five species (one endemic) in China.

1a.       Lamina pinnatifid; sori oblong ........................................................................  1. G. oyamense

1b.       Lamina 2- or 3-pinnate; sori smaller, orbicular.

2a.       Rachis glandular abaxially.

3a.       Stipe sparsely glandular, rachis base and costa base of basal 1–3 pairs of pinnae glandular; veins often forked .......................................................................................................  2. G. jessoense

3b.       Stipe apex, rachis, and costae densely glandular abaxially, other parts also glandular; veins usually simple                                                                                                                3. G. robertianum

2b.       Rachis glabrate, eglandulose.

4a.       Lamina ovate-pentagonal or ternate, lowest pinnae nearly as large as rest of lamina, 2-pinnate-pinnatifid; basiscopic basal pinnules of lowest pinna nearly as large as third pinnae; ultimate pinnules oblong, ± pinnatifid, or shallowly lobed ....................................................................  4. G. dryopteris

4b.       Lamina deltoid-ovate, lowest pinnae smaller than rest of lamina, 3-pinnate-pinnatifid; basiscopic basal pinnules of lowest pinnae nearly as large as fourth pinnae; ultimate pinnules narrowly oblong, usually entire .............................................................................................  5. G. remotepinnatum

1. Gymnocarpium oyamense (Baker) Ching, Contrib. Biol. Lab. Sci. Soc. China, Bot. Ser. 9: 40. 1933.

东亚羽节蕨  dong ya yu jie jue

Polypodium oyamense Baker, J. Bot. 15: 366. 1877; Aspidium krameri (Franchet & Savatier) Christ; Carpogymnia oyamensis (Baker) Α. Lφve & D. Lφve; Currania gracilipes Copeland; C. oyamensis (Baker) Copeland; Dryopteris gymnogrammoides (Baker) C. Christensen; D. oyamensis (Baker) C. Christensen; Gymnocarpium gracilipes (Copeland) Ching; G. oyamense var. gracilipes (Copeland) W. C. Shieh; Nephrodium gymnogrammoides (Baker) Diels; N. krameri (Franchet & Savatier) Diels; Phegopteris krameri (Franchet & Savatier) Makino; P. oyamensis (Baker) Alderwerelt; Polypodium gymnogrammoides Baker; P. krameri Franchet & Savatier; P. krameri var. incisum Franchet & Savatier.

Rhizomes long creeping, 1.5–2(–3) mm in diam., clothed with red-brown broadly lanceolate scales, glabrate when old. Fronds distant; fertile fronds (13–)25–45(–50) cm; stipe stramineous, shiny, (7–)12–25(–31) cm, 1.5–2 mm in diam., terete abaxially and grooved adaxially, scaly at base, upward glabrate, articulate to lamina at apex; lamina ovate-triangular, (7–)10–18(–22) Χ (4–)6–13(–20) cm, herbaceous, glabrate, base cordate, pinnatipartite 4–5 mm to rachis, apex acuminate; pinnae 6–9(–13) pairs, opposite, spreading, approximate, broadly lanceolate, falcate, base decurrent to narrow rachis wing, shallowly lobed to pinnatipartite, apex acute or acuminate; basal pair of pinnae descending, often broadly lanceolate, (2–)4–7(–10) Χ 1–1.8(–3) cm; second pair slightly longer than basal pinnae or subequal, spreading, shallowly lobed; pinna lobes obtuse-rounded, entire or crenate; veins pinnate, lateral veins (2–)4 or 5(–8) pairs, simple, slightly visible; rachis inserted obliquely and articulate to stipe; upper part of stipe, lower part of rachis, and bases of costae in lower pinnae sparsely pubescent with short glands, or glabrate. Sori oblong, medial on veins, biseriate along two sides of costules, exindusiate. Spore-wall surface rugate, foveolate. 2n = 80, 160.

Damp areas in forests, in mosses on rocks; 300–2900 m. Anhui, Chongqing, Gansu, Guizhou, Henan, Hubei, Hunan, Jiangxi, Shaanxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [NE India, Japan, Nepal, New Guinea, Philippines].

2. Gymnocarpium jessoense (Koidzumi) Koidzumi, Acta Phytotax. Geobot. 5: 40. 1936.

羽节蕨  yu jie jue

Dryopteris jessoensis Koidzumi, Bot. Mag. (Tokyo) 38: 104. 1924; Aspidium dryopteris (Linnaeus) Baumgarten var. longulum Christ; Carpogymnia disjuncta (Ruprecht) Α. Lφve & D. Lφve; C. jessoensis (Koidzumi) Α. Lφve & D. Lφve; Dryopteris linnaeana C. Christensen; D. linnaeana var. jessoensis (Koidzumi) C. Christensen; Gymnocarpium dryopteris (Linnaeus) Newman subsp. disjunctum (Ruprecht) Sarvela; G. dryopteris var. disjunctum (Ruprecht) Ching; G. longulum (Christ) Kitagawa; G. remotum (Hayata) Ching, p.p.; G. robertianum (Hoffmann) Newman subsp. longulum (Christ) Toyokuni; G. robertianum var. longulum (Christ) H. Itτ ex Nakai et al.; Lastrea jessoensis (Koidzumi) Akasawa; L. robertiana var. longula (Christ) Ohwi; Polypodium dryopteris Linnaeus var. disjunctum Ruprecht.

Rhizomes long creeping, apex clothed with brownish ovate-lanceolate scales; fronds distant, sometimes approximate. Fertile fronds (16–)20–50(–76) cm; stipe stramineous, (8–)15–32(–51) cm, up to 3.5 mm in diam., base sparsely scaly, upper part glabrate; lamina pinnate-pinnatifid or 2-pinnate-pinnatipartite, deltoid-ovate, (7–)15–20(–27) Χ (7–)14–22(–30) cm, herbaceous or papery, base rounded, apex acuminate; pinnae (3–)5–8 pairs, opposite, oblique, basal 1–4 pairs stalked, articulate to rachis; basal pinnae largest, narrowly triangular, (4–)8–15(–18) Χ 3–7(–11) cm at base, base subtruncate, with stalk (0.8–)1–2.5(–3.5) cm, slightly oblique, pinnate-pinnatifid or pinnate-pinnatipartite, apex acuminate; pinnules 5–8 pairs, deltoid-lanceolate, base subtruncate, apex acuminate, opposite or subopposite; basal one to several pairs articulate to costa, usually sessile, sometimes basal pair shortly stalked, 1–3(–12) mm; basal basiscopic pinnules longest, 1–5(–7) Χ (0.7–)1–2.3 cm, descending; pinnule segments 5–10 pairs, oblong to narrowly ovate, base free or adnate to narrow wing, margin crenate, apex rounded-obtuse; second basal pair of pinnae (2–)4–5(–7.5) cm apart from basal pair, narrowly triangular, much smaller than lowest pinnae, 4–8(–12) cm; third and upper pairs of pinnae broadly lanceolate, gradually smaller; veins pinnate in segment, veins usually forked, sometimes simple, very oblique, visible; stipe apex, rachis, and costae with hyaline or pale yellow short glands. Sori small, orbicular or oblong, abaxial on veins, submarginal, exindusiate. Spore wall surface rugate, foveolate. 2n = 80.

Wet areas in forests, mountain slopes; 400–4000 m. Gansu, Guizhou, Hebei, Heilongjiang, Henan, Jilin, Liaoning, Nei Mongol, Ningxia, Qinghai, Shaanxi, Shanxi, Sichuan, Xinjiang, SE Xizang, NW Yunnan [Afghanistan, Bhutan, N India, Japan, Korea, Nepal, N Pakistan, E Russia; NW North America].

3. Gymnocarpium robertianum (Hoffmann) Newman, Phytologist 4: 371, append. 24. 1851.

密腺羽节蕨  mi xian yu jie jue

Polypodium robertianum Hoffmann, Deutschl. Fl. 2: 20[??add. et emend. 10]. 1795; Aspidium calcareum (J. Smith) Baumgarten; A. robertianum (Hoffmann) Luerssen; Carpogymnia robertiana (Hoffmann) Α. Lφve & D. Lφve; Dryopteris robertiana (Hoffmann) C. Christensen; Gymnocarpium altaycum C. Y. Yang; Phegopteris calcarea (J. Smith) Fιe; Polypodium calcareum J. Smith; P. dryopteris var. calcareum (J. Smith) A. Gray; P. dryopteris var. glandulosum Neilet; P. dryopteris var. robertianum (Hoffmann) Hooker & Baker[??auth.]; P. robertianum (Hoffmann) Fιe[??auth.].

Rhizomes long creeping, apex clothed with brownish ovate-lanceolate scales. Fronds distant; fertile fronds 18–40 cm; stipe stramineous, 11–25 cm, 0.5–1 mm in diam., base sparsely scaly, upper part glabrate; lamina 2-pinnate to 3-pinnatifid, deltoid-ovate, 7–13 Χ 4–8 cm, herbaceous or papery, base rounded, apex acuminate; pinnae 5–8 pairs, opposite, oblique, basal 3 or 4 pairs stalked, articulate to rachis, basal pair largest, narrowly triangular, 4.5–8.5 Χ 3–4.5 cm at base, base subtruncate, with stalk 6–13 cm, slightly oblique, pinnate, apex acuminate; pinnules 4–6 pairs, opposite or subopposite, deltoid-lanceolate or narrowly oblong, base broadly cuneate, basal 1 or 2 pairs articulate to costae, subsessile, apex acuminate; basal basiscopic pinnules longest, descending, 1.5–2.7 cm Χ 6–8 mm, pinnate or pinnatipartite; pinnule segments 5–7 pairs, oblong to narrowly ovate, separate or adnate[??connate] at base, crenate, apex rounded-obtuse; second basal pair of pinnae 1.7–3.3 cm apart from basal pinnae, narrowly triangular, much smaller than basal pinnae, 2–5.5 cm; third and upper pairs of pinnae broadly lanceolate, gradually smaller; veins pinnate in segment, simple, sometimes forked, oblique, visible; stipe (at least upper part), lower part of rachis, costae, and midribs pubescent with short glands. Sori small, orbicular or oblong, abaxial on veins, submarginal, exindusiate. 2n = 160.

Shaded places in forests, mountain slopes; 1500–2500 m. Qinghai, Xinjiang [Pakistan; widespread in Europe and North America].

4. Gymnocarpium dryopteris (Linnaeus) Newman, Phytologist 4: 371 [append. xxiv]. 1851.

欧洲羽节蕨  ou zhou yu jie jue

Polypodium dryopteris Linnaeus, Sp. Pl. 2: 1093. 1753; Aspidium dryopteris (Linnaeus) Baumgarten; Carpogymnia dryopteris (Linnaeus) Α. Lφve & D. Lφve; Dryopteris linnaeana (Linnaeus) C. Christensen; Lastrea dryopteris (Linnaeus) Bory.

Rhizomes slender, creeping, black, shiny, apex clothed with brown ovate-lanceolate scales. Fronds distant; fertile lamina (15–)20–30(–50) cm; stipe stramineous, 10–22(–35) cm, slender, with sparse scales at base; lamina nearly tripartite, usually 2-pinnate-pinnatifid, pentagonal-ovate or broadly ovate-triangular, 7–15(–20) cm long and wide, thinly herbaceous or submembranous, base broadly cuneate, apex acuminate; basal pair of pinnae nearly as large as others portion of lamina, narrowly triangular, (3.5–)5–9(–12) Χ 2.5–4(–7) cm, pinnate-pinnatifid, base subtruncate, with stalk (0.8–)1–1.5(–2.5) cm, portion of rachis between basal pinnae and central lamina ca. 3 cm; pinnules 5 or 6 pairs, oblong-lanceolate, 1.5–2(–4) Χ 0.5–2 cm, base rounded-cuneate, sessile, apex acute or acuminate, opposite or subopposite, spreading; largest pinnules with 6–10 pairs of segments, segments approximate, oblong to narrowly ovate, ca. 4 mm, lobed to narrow costular wing, entire to shallowly lobed at margin, rounded-obtuse at apex; second basal pair of pinnae 1.5–4 cm apart from basal pair, sometimes shortly stalked, upper pinnae sessile; veins pinnate in segment, simple, oblique, visible abaxially; rachis and costae slender, eglandular. Sori small, exindusiate, orbicular, abaxial on veins. Spore wall surface rugate, foveolate. 2n = 160.

Damp areas in coniferous forests; 300–2900 m. Heilongjiang, Jilin, Liaoning, Nei Mongol, Shaanxi, Shanxi, Xinjiang [Japan, Korea; Europe, North America].

American Gymnocarpium dryopteris is an allotetraploid species that arose following polyploidization and hybridization between G. disjunctum (Ruprecht) Α. Lφve & D. Lφve and G. appalachianum Pryer & Haufler.

Sarvela (1976) noted that a specimen of Gymnocarpium dryopteris from Japan was indusiate; the indusia were reniform, ca. 0.5 mm in diam., hyaline, glabrate, erose at margin. It may be an example of atavism.

5. Gymnocarpium remotepinnatum (Hayata) Ching, Bull. Chin. Bot. Soc. 1(2): 14. 1935 [??Hu & Ching, Icon. Fil. Sin. 4: t. 127. 1937].

细裂羽节蕨  xi lie yu jie jue

Dryopteris remotepinnata Hayata, Suppl. Icon. Pl. Formosan. 6: 108. 1917; Gymnocarpium remotum (Hayata) Ching, p.p. [basionym is nom. superfl.]; Thelypteris remote-pinnata (Hayata) Alston.

Rhizomes slender, creeping, with sparse brownish ovate-lanceolate scales. Fronds distant; fertile fronds 20–30 cm; stipe stramineous, (10–)12–20 cm, slender, fragile, sparsely scaly at base; lamina 3-pinnate or 3-pinnate-pinnatifid, triangular, 10–14 Χ 6–9 cm, thinly herbaceous, base subtruncate, apex acuminate; pinnae 5 or 6 pairs, opposite, basal 2 pairs shortly stalked, upper pinnae sessile, basal pinnae largest, narrowly triangular, 6–8 Χ 5–6 cm, base subtruncate, stalk 1–2.2 cm, articulate to rachis, 2-pinnate, apex acuminate, spreading; pinnules 5 or 6 pairs, opposite, oblong-lanceolate, 1.5–2 Χ ca. 1 cm, base broadly rounded, sessile, apex acuminate, spreading; pinnule segments or secondary pinnules narrowly oblong, separate or lobed to costular wings, entire but lobed on basiscopic side, apex rounded-obtuse; second basal pair of pinnae 2–3 cm apart from basal pinnae, smaller, ascending, oblong, falcate, base rounded and equilateral, sessile; upper pinnae gradually smaller than second basal pinnae; veins pinnate in segment, simple, very oblique, visible abaxially; lamina glabrate, abaxial surface of rachis and costae eglandular. Sori small, brown, orbicular, exindusiate, abaxial on veins. 2n = 80*.

* Coniferous forests, on rocks at forest margins; 2500–3400 m. Taiwan, NW Yunnan.

2. CYSTOATHYRIUM Ching, Acta Phytotax. Sin. 11: 22. 1966.

光叶蕨属  guang ye jue shu

Wang Zhongren (王中仁); Masahiro Kato

Athyrium Kato & Kramer, p.p.[??ok, not Roth (1799)]

Plants evergreen, medium-sized. Rhizomes short, ascending, bearing remaining stipe bases and dense thick roots, clothed with dark brown ovate-lanceolate scales at apex. Fronds approximate; fertile fronds 40–45 cm; stipe brown at base, upward stramineous, up to 7–8 cm, base ca. 2 mm in diam., slightly swollen, with 1 or 2 lanceolate scales, upper part glabrate, grooved adaxially; rachis grooved adaxially, glabrate; lamina pinnate-pinnatifid, pale green, narrowly lanceolate, up to 35 Χ 6–8 cm at middle, subpapery, glabrate, gradually narrowed toward both ends, with scattered orange glands on abaxial surface; pinnae pinnatipartite, up to 30 pairs, subopposite, spreading, sessile, ca. 1 cm apart (lower pinnae more widely apart), longest middle pinnae 3–4 Χ ca. 1 cm at base, narrowly falcate-lanceolate, base inequilateral (acroscopic side broader, truncate, basiscopic side narrowly cuneate or obtuse-rounded), pinnatipartite, lobed 4/5 or close to costae, apex acuminate and falcate; lower pinnae gradually shorter than upper pinnae, basal pair shortest, ca. 1 cm, triangular; pinna lobes up to ca. 10 pairs, oblique, oblong, obtuse at apex, separated by narrow incisions, acroscopic basal lobe slightly longer than basiscopic one, basal 2 lobes largest, 5–8 Χ ca. 3 mm, upper lobes gradually shorter, basal basiscopic lobe almost broadly ovate, slightly shorter, entire at margin, or basal 1 or 2 pairs slightly crenate; costae grooved adaxially; small processes of unknown nature one to few, on abaxial surface of lateral veins in most middle pinna apices. Veins per lobes pinnate, visible, anadromous, veinlets 3–5 pairs, simple but basal veins frequently forked, oblique, reaching margin of lobes. Sori orbicular, single per pinna lobe, abaxial on basal acroscopic veins, close to costae; indusia pale green, broadly ovate or orbicular, thinly membranous, fugacious, basiscopic to receptacle, inferior (i.e., hidden by sporangia), partly covering sori when young, hidden by sporangia at maturity, persistent; annulus consisting of 12 or 13 thickly walled cells. Spores dark brown, bean-shaped, darkened, perispore with dense conical spines.

Monotypic genus: the one species endemic to China.

Cystoathyrium is similar to Athyrium and Cystopteris. It differs from Athyrium by the rhizome shortly ascending, glabrate, the stipe base not beak-shaped, spore surface with conical spines, the indusia basiscopic and inferior. It also differs from Cystopteris by the stipe short compared to the lamina, the lamina bipinnatifid, gradually narrowed toward the base, almost papery and evergreen, and the pinnae numerous, lanceolate and falcate.

1. Cystoathyrium chinense Ching, Acta Phytotax. Sin. 11: 23. 1966.

光叶蕨  guang ye jue

Description?

* Damp areas of forest understories; 2400–2500 m. W Sichuan.

The type of Cystoathyrium chinense is from Sichuan (Erlang Shan, Tianquan). Because it has not been collected recently, it is possible that it is now extinct.

3. ACYSTOPTERIS Nakai, Bot. Mag. (Tokyo) 47: 180. 1933.

亮毛蕨属  liang mao jue shu

Wang Zhongren (王中仁); Masahiro Kato

Cystopteris Luerssen (1883), non Bernhardi (1806); Cystopteris subg. Acystopteris (Nakai) Blasdell.

Plants terrestrial, medium-sized. Rhizome creeping, forked, sparsely scaly; scales shiny, lanceolate or ovate-lanceolate, margin with sparse glandular-hairlike teeth, apex long acuminate. Fronds approximate; stipe chestnut-brown or stramineous, nearly as long as lamina, with scales, hyaline multicellular hairs, and scalelike hairs, grooved adaxially; lamina 2-pinnate to 3-pinnate-pinnatifid, broadly ovate to ovate-lanceolate, thinly herbaceous, not narrowed to base, apex subacute or acuminate; pinnae mostly subopposite, spreading or ascending, pinnatisect to 2-pinnate, broadly lanceolate to lanceolate, slightly falcate at apex, equilateral, truncate or rounded-cuneate at base, subsessile or very shortly stalked, apex acuminate; basal pair of pinnae not shortened; pinnules sessile, alternate, only basal pair of pinnules subopposite, slightly shorter than upper pinnules, ovate-lanceolate or lanceolate, equilateral, base subtruncate or rounded-cuneate, basiscopic base slightly wider than acroscopic base, apex obtuse or acuminate; pinnule lobes oblong, sessile or adnate to costae or costules, lobed or parted, obtuse. Veins visible but not clearly so, free, veins pinnate, simple or forked, reaching marginal teeth of lobes. Lamina surface, all costae, and veins with hyaline long multicellular hairs and narrow scalelike hairs. Sori small, orbicular, abaxial on veins, sorophore protuberant; sporangia with slender stalks, annulus cells 12–14; indusia small, laterally equilateral, pale green, broadly ovate, membranous, glandular hairy, sparsely ciliate at margin, attached proximally to sorophore, hidden by sporangia at maturity; spores yellow, bean-shaped or narrowly spheroidal, perispore with dense inflated, bulbous processes. x = 42.

Three species: tropical to temperate areas of SE and E Asia and New Zealand; three species (one endemic) in China.

1a.       Plants usually less than 90 cm; orange unicellular glands absent on lamina and at indusium margin; stipe, rachis, and costae chestnut-black or purple-brown
                                                                                                                                                             1. A. japonica

1b.       Plants to 1 m or longer; orange unicellular glands present on abaxial surface of lamina and at indusium margin; stipe and rachis stramineous or brownish red, costae stramineous.

2a.       Stipe and rachis stramineous; multicellular hairs dense ..................................  2. A. tenuisecta

2b.       Stipe and rachis brownish or dull-brown; multicellular hairs sparse ..............  3. A. taiwaniana

1. Acystopteris japonica (Luerssen) Nakai, Bot. Mag. (Tokyo) 47: 180. 1933.

亮毛蕨  liang mao jue

Cystopteris japonica Luerssen in Engler, Bot. Jahrb. Syst. 4: 363. 1883.

Plants summer-green. Rhizome creeping, 2–4 mm in diam., with sparse yellowish brown, broadly lanceolate, thinly membranous scales. Fronds approximate; fertile frond (12–)40–60(–90) cm; stipe chestnut-black or purple-brown, (6–)15–25(–40) cm, (1–)2–3(–4) mm in diam., grooved adaxially, with sparse scales at base, upward glabrate, shiny; lamina 2- or 3-pinnate, broadly ovate to deltoid-ovate, (6–)20–35(–50) Χ (3.5–)15–18(–40) cm, not narrowed to base, herbaceous, base subcordate, apex shortly acuminate or acuminate; pinnae 10–15 pairs, opposite or subopposite, 3–4 cm apart, spreading or ascending, subsessile or very shortly stalked, basal pair not shorter, oblong or broadly lanceolate; upper pinnae lanceolate, (1.8–)8–15(–25) Χ (1–)3–6(–11) cm, base subtruncate, apex acuminate; pinnules 10–24 pairs, sessile, alternate, subequilateral, basiscopic pinnules slightly longer than acroscopic pinnules; basal pair slightly shorter, subopposite or anadromous, oblong, (0.5–)2.5–4(–6) Χ (3–)6–10(–20) mm, costae stramineous or only base chestnut-brown, base truncate or broadly cuneate, apex obtuse; pinnule segments 5–14 pairs, spreading or slightly ascending, oblong, crenate, apex obtuse; veins pinnate in segment, simple, reaching marginal teeth. Hyaline multicellular long hairs and scalelike hairs sparse on both sides of veins, costae, rachis, and stipe. Sori orbicular, abaxial on basal acroscopic veins; indusia pale green, broadly ovate, small, membranous, glandular, sparsely ciliate at margin, proximal to sorophore, hidden by sporangia at maturity, persistent. Spores yellow, bean-shaped or long spheroidal, perispore with dense inflated, bulbous processes. 2n = 84*, 168*.

Forests in valleys; 400–2800 m. Chongqing, Fujian, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Taiwan, Yunnan, Zhejiang [Japan].

2. Acystopteris tenuisecta (Blume) Tagawa, Acta Phytotax. Geobot. 7(2): 73. 1938.

禾秆亮毛蕨  he gan liang mao jue

Aspidium tenuisectum Blume, Enum. Pl. Javae 2: 170. 1828; Alsophila tenuisecta Blume ex T. Moore; Asplenium tenuisectum (Blume) Hooker; Athyrium tenuisectum (Blume) T. Moore; Cornopteris tenuisecta (Blume) Tardieu; Cystopteris formosana Hayata; C. setosa Beddome; C. tenuisecta (Blume) Mettenius; Davallia setosa (Beddome) Baker (1874), not J. Smith (1808); Lastrea setosa Beddome (1868), not C. Presl (1851), nor T. Moore (1858).

Rhizomes creeping, 2–5 mm in diam., with sparse brownish lanceolate thinly membranous scales. Fronds approximate; fertile frond (22–)65–80(–150) cm; stipe pale stramineous, (7–)30–40(–90) cm, (1.5–)3–4(–5) mm in diam. at base, grooved adaxially, clothed with brownish lanceolate scales and sparse multicellular hairs at base; lamina 3-pinnate to 3-pinnate, ovate to ovate-lanceolate, (14–)35–50(–70) Χ (7–)20–25(–60) cm, herbaceous, base subcordate, apex acuminate; pinnae 15–20 pairs, subopposite, subsessile or shortly stalked, broadly lanceolate to lanceolate, acuminate, slightly falcate, equilateral, base subtruncate; basal 1 or 2 pairs broadly lanceolate, (3.5–)15–20(–33) Χ (2–)4–8(–20) cm, base subtruncate or rounded-cuneate, apex long acuminate; pinnules 10–25 pairs, basal pair slightly shorter, acroscopic pinnule slightly shorter than basiscopic, broadly lanceolate, (1–)2.5–5(–10) Χ (0.4–)1–2(–3) cm, base truncate, sessile, apex obtuse or acuminate; pinnule segments oblong, 4–10 Χ 2–3 mm, sessile or ± adnate, lobed, obtuse; veins pinnate in segment, simple or forked. Orange or yellow small glands borne on abaxial surface and at indusium margin, hyaline multicellular long hairs present on veins, costules, costae, rachis, and stipe, and subulate scales adaxially. Sori orbicular, abaxial on basal acroscopic veins; indusia small, pale green, broadly ovate, membranous, proximal to sorophore, hidden by sori at maturity, persistent. Spores yellow, bean-shaped or narrowly spheroidal, perispore with dense inflated, bulbous processes. n = 41, 42, 84.

Forests in valleys and damp streamsides; 700–2600 m. Guangxi, Sichuan, Taiwan, Xizang, Yunnan [Bhutan, NE India, Indonesia, S Japan, Malaysia, Myanmar, Nepal, New Zealand, Philippines, Singapore, Thailand, Vietnam].

3. Acystopteris taiwaniana (Tagawa) Α. Lφve & D. Lφve, Taxon 26: 326. 1977 [and in Lφve, Lφve & Pic. Ser., Cytotax. Atlas Pterid. 269. 1977].

台湾亮毛蕨  tai wan liang mao jue

Cystopteris japonica var. taiwaniana Tagawa, Acta Phytotax. Geobot. 4: 56[55, 57]. 1935; Acystopteris japonica var. taiwaniana (Tagawa) W. C. Shieh.

Intermediate between Acystopteris japonica and A. tenuisecta. Stipe pale brown-red, costae stramineous, lamina with pale yellow glandular cells on abaxial surface. Spores normal; diploid or tetraploid. n = 41. [??make parallel]

* 1600–2600 m. Taiwan.

4. CYSTOPTERIS Bernhardi in Schrader, Neues J. Bot. 1(2): 26. 1805, nom. cons.

冷蕨属  leng jue shu

Wang Zhongren (王中仁); Chris Haufler

Cystea J. Smith; Filix Ludwig (1757), not Sιguier (1754); Rhizomatopteris A. P. Khokhrjakov.

Plants small, summer-green. Rhizomes long or shortly creeping, blackish brown, glabrate or densely red-brown pubescent, sparsely scaly; scales brown or brownish, thin, ovate to broadly lanceolate. Fronds distant, approximate or caespitose, thin; stipe dark brown at base, upper part stramineous or chestnut-colored; lamina 2- or 3-pinnate, rarely 4-pinnate or more divided, ovate-lanceolate, ovate-triangular, or subpentagonal; pinnae with anadromous pinnules, shortly stalked, unequally sided or subequilateral at base; minutely toothed at segment margin; veins free, forked or pinnate, terminating in teeth or emarginations. Lamina thinly herbaceous or herbaceous, green when dried, stipe, rachis, costae, and costules with few small scales at base, and multicellular hairs or unicellular glandular hairs, lamina usually glabrate. Sori orbicular, abaxial on veins, sorophore slightly protuberant, annulus of sporangium consisting of 14–16 cells; indusia ovate, lanceolate, orbicular, or shallow cup-shaped, attached at proximal side of sorophore, membranous, persistent, covering sporangia when young, hidden at maturity. Spores bilateral, dark brown, bean-shaped, perispore usually with spines, rarely with folds. x = 42.

More than 20 species and many hybrids worldwide: mainly in the temperate and frigid-temperate zones, and tropical mountains; 11 species (six endemic) in China.

1a.       Rhizomes shortly creeping or ascending; fronds approximate or caespitose; lamina lanceolate to broadly lanceolate; veins terminating in teeth, rarely in emarginations.

2a.       Veins terminating in teeth.

3a.       Perispore echinate; on calcareous soil ............................................................  1. C. fragilis

3b.       Perispore rugose or verrucose, not echinate; on acidic soil .........................  2. C. dickieana

2b.       Veins terminating in emarginations.

4a.       Plant usually less than 10 cm tall; stipe slender, slightly longer than lamina or nearly as long as lamina, chestnut-colored .......................................................................................  3. C. kansuana

4b.       Plant up to 29 cm tall; stipe thick (ca. 1 mm), shorter than lamina, dark purple at base .  4. C. guizhouensis

1b.       Rhizomes long creeping with fronds distant; lamina broadly ovate, ovate-triangular, or subpentagonal; veins terminating in emarginations.

5a.       Lamina subpentagonal with basal basiscopic pinnules of basal pinnae prominently elongated.

6a.       Ultimate segments not revolute at margin; spore surface shortly spinous or verrucose  5. C. montana

6b.       Ultimate segments revolute at margin; spore surface rugately folded ..............  6. C. modesta

5b.       Lamina broadly ovate or ovate-triangular with basal basiscopic pinnules of basal pinnae not elongated.

7a.       Indusia glandular; plant slender ....................................................................  7. C. sudetica

7b.       Indusia not glandular, plant moderately stout.

8a.       Lamina with short glandular hairs ................................................................  8. C. tibetica

8b.       Lamina lacking short glandular hairs.

9a.       Fronds up to 50 cm or longer, lamina to 4-pinnatisect, perispore densely echinate  9. C. deqinensis

9b.       Fronds less than 50 cm, lamina 2- or 3-pinnate, perispore sparsely echinate.

10a.     Ultimate segments 3–5 mm wide, indusia attached to ca. 1/2 of circumference of receptacle        10. C. moupperensis

10b.     Ultimate pinnules or segments 5–7 mm wide, indusia attached to ca. 2/3 of circumference of receptacle .........................................................................................  11. C. pellucida

1. Cystopteris fragilis (Linnaeus) Bernhardi in Schrader, Neues J. Bot. 1(2): 26. 1805.

冷蕨  leng jue

Polypodium fragile Linnaeus, Sp. Pl. 2: 1091. 1753; Aspidium fragile (Linnaeus) Swartz; Athyrium fragile (Linnaeus) Sprague; Cyclopteris fragilis (Linnaeus) Gray; Cystea fragilis (Linnaeus) J. Smith; Cystopteris filix-fragilis (Linnaeus) Gilibert.

Rhizomes shortly or long creeping, scaly at apex and stipe bases; scales brownish, broadly lanceolate. Fronds approximate or caespitose; fertile fronds (3.5–)20–35(–49) cm; stipe brown at base, upper part stramineous or chestnut-colored, usually shorter than lamina, nearly as 1/3–2/3 as long as lamina, occasionally slender and slightly longer than lamina when plant in crevices, 5–14(–20) cm, (0.2–)1–1.5 mm in diam., with sparse, shiny scales; lamina usually 2-pinnate to 2-pinnate-pinnatifid, occasionally pinnate or 3-pinnate, lanceolate to broadly lanceolate, 17–28 Χ (0.8–)4–5(–8) cm, shortly acuminate at apex; pinnae 12–15 pairs, ascending, basal 1 or 2 pairs slightly shortened, or nearly not shortened, ovate to ovate-lanceolate, (0.4–)2–4(–7) Χ (0.2–)1–2.5 cm[??], base parallel to rachis on acroscopic side, ± cuneate on basiscopic side, obtuse or shortly acuminate, toothed at apex, subopposite, subsessile, widely separated from each other, usually 1–2 Χ as long as wide, 1.5–4.5 cm[??]; pinnules 5–7 pairs, ovate or oblong, acroscopic side truncate, basiscopic side cuneate at base, sessile or shortly stalked, entire or toothed at margin, or pinnatifid, rounded or obtuse and toothed at apex; middle pinnae similar to lower pinnae, but slightly longer, separated from each other by 1.2–2.5 cm, subopposite or alternate, subsessile; upper pinnae pinnatipartite, toothed only on apices and acroscopic margin; veins pinnate, costules slightly tortuous, terminating in entire teeth. Lamina herbaceous when dried, green or yellow-green. Rachis and costae, especially their bases, ± with sparse unicellular to long multicellular hairs, even with few scale hairs. Sori small, orbicular, medial on veins, 2–4 pairs per pinnule, 1 or 2 sori at acroscopic side in upper pinnules; indusia pale green or brownish, ovate to lanceolate, membranous. Spores dark brown, perispore surface with regular, denser spines. 2n = 84*, 168, 252, 336.

Under alpine shrubs, rock crevices on shaded slopes, foot of rocks, damp areas at streamsides; (200–)1500–4500(–4800) m. Anhui, Gansu, Hebei, Heilongjiang, Henan, Jilin, Liaoning, Nei Mongol, Ningxia, Qinghai, Shaanxi, Shandong, Shanxi, W Sichuan, Taiwan, Xinjiang, Xizang, Yunnan [Afghanistan, India, Japan, Kashmir, Korea, Mongolia, Nepal, Pakistan, Russia; Africa, SW Asia (Iran, Turkey), Europe, N and C mountains and Himalaya, North America, South America].

The type is from Europe.

2. Cystopteris dickieana R. Sim, Gard. Farmer’s J. 2: 308. 1848.

皱孢冷蕨  zhou bao leng jue

Cystopteris fragilis var. dickieana (R. Sim) Lindberg in Medd. Soc. Fauna & Fl. Fenni 32: 21–24. 1905[(R. Sim) T. Moore, Index Fil. 282. 1861]; C. baenitzii Dφrfler; C. fragilis subsp. dickieana (R. Sim) Hylander; C. sikkimensis Ching ex Bir.

Very similar to Cystopteris fragilis, hardly distinguished from it morphologically except in spore ornamentation, i.e., perispore rugose or verrucose not echinate. It is reported that they can make sterile hybrids and form a species complex with other species. 2n = 168, 252. [??make parallel]

Sympatric in the distribution area of Cystopteris fragilis, valleys, rock crevices on mountain slopes, on rocks of forest understories, damp areas in grasslands; 1400–5400(–5600) m. Gansu, Hebei, Qinghai, Shaanxi, Sichuan, Taiwan, Xinjiang (common), Xizang (common), Yunnan [Afghanistan, India, Nepal, Pakistan; Africa, Europe, North America].

The taxonomic status of Chinese Cystopteris dickieana is uncertain. Scottish and American C. dickieana are considered conspecific with C. fragilis, based on the variable echinate and rugose spore ornamentation in C. fragilis.

3. Cystopteris kansuana C. Christensen, J. Wash. Acad. Sci. 17: 499. 1927.

西宁冷蕨  xi ning leng jue

Cystopteris Χ kansuana Blasdell.

Rhizomes slender, shortly creeping, sparsely scaly. Fronds approximate; fertile fronds 10–19 cm; stipe chestnut-colored, 6–11 cm, less than 1 mm in diam., base with few brown lanceolate scales, upper part glabrate; lamina bipinnatifid, pale green or dark brown when dried, lanceolate, 4–10 Χ 1.3–3 cm, thinly herbaceous, base obtuse, apex long acuminate; pinnae 5–8 pairs, basal pinnae 1–2.4 cm apart from, and slightly shorter than, second basal pinnae, subopposite; upper pinnae alternate, ascending, subsessile or shortly stalked (stalk 1–2 mm); lower pinnae ovate to narrowly ovate, 9–20 Χ 4–8 mm, acroscopic base broadly cuneate, basiscopic base narrowly cuneate, lobed to narrow costal wings, apex acuminate; pinna segments ca. 3 pairs, ascending; basal acroscopic segment largest, narrowly ovate, 4–8 Χ 2–3 mm, crenate or dentate-pinnatipartite, apex acute or obtuse-rounded; upper segments oblong or linear-oblong, acuminate, with 2 or 3 small teeth, entire at margin. Veins in basal acroscopic lateral segment pinnate, upper or basiscopic lateral veins simple or occasionally forked, terminating in laminar emarginations. Sori yellow-brown, orbicular, medial on veins; indusia pale yellow-brown, ovate, entire at margin, persistent. Spore with echinate perispore.

* Shaded rock crevices; 3000–4500 m. Gansu, Qinghai (type from Xining), Sichuan, Xizang, Yunnan.

4. Cystopteris guizhouensis X. Y. Wang & P. S. Wang, Acta Bot. Yunnan. 19(2): 141. 1997.

贵州冷蕨  gui zhou leng jue

Rhizomes shortly creeping, scaly at apex along with stipe bases; scales brown, ovate-lanceolate. Fronds approximate or caespitose, up to 29 cm; stipe base dark purple, upper part stramineous, 7–9 cm, nearly 1/2 as long as lamina; lamina bipinnatifid, lanceolate, 18–21 Χ 4–5 cm at middle, thinly herbaceous when dried, dark green, glabrate on surfaces, base slightly narrowed, apex long acuminate; pinnae 12–15 pairs, spreading, basal 1 or 2 pairs slightly shorter than upper pinnae, subopposite, sessile, oblong or ovate-oblong, 2.5–2.8 Χ 1–1.2 cm, subequilateral, cuneate at base, obtuse at apex; pinna segments 8–10 pairs, often oblong, crenate, apex rounded or truncate; veins free, terminating in emarginations. Sori small, orbicular, usually in one row on either side of costa; indusia broadly ovate, membranous, persistent. Perispore echinate, spines 6–7 μm.

* Rock crevices in forests; ca. 2800 m. Guizhou.

5. Cystopteris montana (Lamarck) Bernhardi ex Desvaux in Schrader, J. Bot. 1(2): 26. 1805.

高山冷蕨  gao shan leng jue

Polypodium montanum Lamarck, Fl. Franc. 1: 23. 1779; Aspidium montanum (Lamarck) Swartz; Athyrium montanum (Lamarck) Rohl ex Sprengel; Rhizomatopteris montana (Lamarck) A. P. Khokhrjakov.

Rhizomes long creeping, blackish brown, glabrate, with sparse brownish ovate membranous scales, more densely scaly at apex. Fronds distant; fertile fronds 20–49 cm; stipe blackish brown at lower part, upper part stramineous or pale chestnut-colored, (6–)14–31 cm, 1–3 Χ as long as lamina, with sparse brownish ovate scales at base, upper part subglabrate with few scales; lamina 3- or 4-pinnate, rarely 4-pinnate-pinnatifid, subpentagonal, (5–)8–15(–20) cm long and wide, thinly herbaceous, apex acuminate; pinnae 4–7(–10) pairs, basal pinnae subopposite (upper pinnae alternate), spreading, shortly stalked (stalk 3–10 mm), ca. 2.5 cm apart from second basal pinnae, largest, deltoid-ovate or triangular, (2.5–)6–11(–13) Χ (2–)4–7 cm, base subtruncate, strongly inequilateral, 2-pinnate with pinnatipartite secondary pinnules, apex acuminate; pinnules (3–)6–8(–10) pairs, anadromous; acroscopic pinnules triangular, (0.4–)1–2 Χ (6–)8–9(–12) mm, base subtruncate, equilateral, basiscopic pinnules prominently larger than acroscopic pinnules, nearly 2 Χ as long, apex acuminate; basal basiscopic pinnules of lowest pinnae largest, oblong-triangular, 2–3 Χ as long as acroscopic pinnules, 3–5 Χ 1.5–2.5 cm, inserted at costa at nearly right angles, sessile or with stalk 1–2 mm, base truncate, apex acuminate; secondary pinnules ca. 6 pairs, alternate, spreading, ovate to oblong, subacute, sessile, often decurrent to costules; acroscopic basal secondary pinnules shorter, deltoid-ovate, 4–9 Χ 3–6 mm, basiscopic pinnules up to 8–16 Χ 6–9 mm; secondary pinnule segments 4 or 5 pairs, subopposite, ascending, base cuneate, lobed to narrow wing of midrib, apex obtuse-rounded; ultimate segments oblique, lobed or slightly incised, rounded-obtuse. Veins pinnate, costules slightly tortuous, simple or forked, terminating in emarginations. Rachis, costae, and costules with unicellular hairs, multicellular hyaline hairs, or short glandular hairs. Sori small, brown, orbicular, abaxial on veins, 3–7 sori per ultimate segment; indusia pale green or yellow-brown, orbicular, thinly membranous. Perispore with short spinous or verrucose processes. 2n = 168.

Alpine areas in mountains, wet areas in forests; 1700–4500 m. Gansu, Hebei, Henan, Nei Mongol, Ningxia, Qinghai, Shaanxi, Shanxi, Sichuan, Taiwan, Xinjiang, Xizang, Yunnan [N India, Japan, Kashmir, Korea, Nepal, E Pakistan, Russia; E Europe, North America].

6. Cystopteris modesta Ching, Bull. Fan Mem. Inst. Biol. Bot. 10: 5. 1940.

卷叶冷蕨  juan ye leng jue

Cystopteris sudetica A. Braun & Milde var. moupinensis (Franchet) Blasdell, p.p.

Rhizomes long creeping, blackish brown, glabrate, with sparse brownish broadly ovate membranous scales, more densely scaly at apex. Fronds distant; fertile fronds 10–30 cm; stipe brown or chestnut-colored, upward stramineous, 6–17 cm, base with sparse brownish ovate scales, upper part with only few scales; lamina 3-pinnate with secondary pinnules pinnatipartite, ovate, 5–13 Χ 4–7 cm, base broadly cuneate, apex acuminate; pinnae 4 or 5 pairs, alternate, ascending, stalk 3–5 mm; basal pinnae largest, narrowly deltoid-ovate, ca. 7 Χ ca. 2.5 cm, base broadly cuneate, apex acuminate; pinnules 5–9 pairs, alternate, very ascending, ca. 8 mm apart, oblong-ovate, base subtruncate, equilateral, shortly stalked, apex acuminate; basal basiscopic pinnules of lowest pinnae longest, up to 3.2 cm; secondary pinnules 4 or 5 pairs, oblong, 3–6 mm, gradually narrowed to apex, base subsessile, not decurrent, apex obtuse; pinnule segments small, 2 or 3 pairs, approximate, broadly ovate, decurrent to narrow costular wing, subentire, revolute at margin, apex rounded-obtuse; second basal pinnae ca. 3 cm apart from basal pinnae. Veins pinnate in segment, free; rachis, costae, and midribs glabrate. Sori orbicular, abaxial on veins; indusia brown, persistent. Perispore foveolate or reticulate with rugate folds.

* Rock crevices at streamsides; ca. 3600 m. NW Yunnan (watershed between Nujiang and Qiujiang).

Cystopteris modesta is very similar to C. montana, but distinct from it in the revolute segments and foveolate or finely reticulate perispore.

7. Cystopteris sudetica A. Brown & Milde, Jahresber. Schles. Ges. Vaterl. Cult. 92. 1855.

欧洲冷蕨  ou zhou leng jue

Cystopteris leucosoria Schur; Rhizomatopteris sudetica (A. Brown & Milde) A. P. Khokhrjakov.

Rhizomes long creeping, 1–2 mm in diam., rhizome and stipe bases clothed with brown short hairs and few pale brown membranous ovate-lanceolate scales, apex more densely scaly. Fronds distant; fertile fronds (15–)20–30 cm; stipe stramineous, shiny, 10–16(–20) cm, thin; lamina 3-pinnate, broadly ovate or ovate-triangular, 9–15(–20) Χ 8–12(–15) cm, thinly herbaceous or herbaceous when dry, acuminate; pinnae 8–12 pairs, ascending, basal pinnae not shortened, oblong or ovate-lanceolate, (3–)4.5–7(–8) Χ 1.8–3 cm at middle, slightly narrowed, inequilateral at base, stalked (stalk 2–3 mm), subopposite, (1–)1.5–2.5(–3) cm apart from second basal pinnae, 2-pinnate, apex acuminate; pinnules 8–12 pairs, anadromous, acroscopic pinnule slightly shorter than basiscopic one or subequal second basiscopic pinnule largest, ovate or ovate-triangular, 1–2 cm Χ 5–8 mm, unequally sided, acroscopic base parallel to costa, basiscopic base cuneate, subsessile or with 1–2 mm stalk, toothed, obtuse or acute at apex, alternate; secondary pinnules or pinnule segments 4 or 5 pairs, broadly ovate to oblong, basal acroscopic one largest, 5–6 Χ 3–4 mm, broadly cuneate, free or adnate to costule by narrow wing at base, margin lobed, rounded-obtuse or subtruncate, serrate at apex; upper pinnules gradually smaller, oblong-obovate, crenate at apex, basiscopic side entire, acroscopic side with 1 or 2 lobes; upper pinnae lanceolate, pinnatifid, segments serrate at apex, toward base entire at margin; veins visible on both surfaces, simple or 1- or 2-forked, terminating in emarginations. Rachis and costae with sparse or dense short glandular hairs and few long multicellular hairs. Sori small, orbicular, 1 or 2 per ultimate segment, often abaxial on suprabasal veins; indusia pale brown or yellow-brown, suborbicular or shallowly cupular, with sparse tiny glandular hairs. Perispore echinate. 2n = 84, 168.

Coniferous and mixed forests; 900–3300 m. Hebei, Heilongjiang, Jilin, Liaoning, Nei Mongol, Shanxi, Xizang, Yunnan [Japan, Korea, E and W Russia; Europe].

8. Cystopteris tibetica Z. R. Wang, Acta Phytotax. Sin. 32(1): 85. 1994.

藏冷蕨  zang leng jue

Rhizomes long creeping, 1.5–2 mm in diam., rhizome and stipe base clothed with brown hairs and few membranous scales. Fronds distant; fertile fronds 15–20 cm; stipe brownish at base, upward stramineous, 6–14 cm, 1–1.5 mm in diam., base clothed with brown hairs and few broadly ovate or ovate-lanceolate scales; lamina 3-pinnate, deltoid-ovate, 6–11 Χ 3–5 cm, acuminate; pinnae 8–10 pairs, alternate, basal pinnae subopposite, 1.2–3 cm apart from second basal pinnae, ascending, largest, oblong or ovate-lanceolate, 3–5 Χ 1.2–1.5 cm, base slightly narrowed and inequilateral, with stalk 1.5–3 mm, apex acuminate; upper pinnae lanceolate; pinnules 6–8 pairs, anadromous, acroscopic one usually slightly shorter than basiscopic one, second basal basiscopic pinnules largest, deltoid-ovate, 6–8 Χ 4–5 mm, inequilateral at base, acroscopic base parallel to costae, basiscopic base broadly cuneate, very shortly stalked, toothed, obtuse at apex, alternate; secondary pinnules or pinnule segments ca. 3 pairs, basal acroscopic largest, ovate, oblong, or obovate, 2–3Χ 1.5–2 cm, base cuneate and adnate to costular wing or partly separate, serrate, apex obtuse; upper segments rhomboid or shortly linear, only acroscopic side and apices toothed; veins visible on surfaces, pinnate, terminating in emarginations. Fronds thinly herbaceous when dried, green, clothed with short glandular hairs on both surfaces. Sori small, orbicular, 1 or 2 sori per ultimate segment, abaxial on suprabasal veins; indusia pale brown, orbicular or shallowly cupular, frequently with few glands, covering sporangia when young, hidden at maturity.

* Alpine slopes, damp areas in coniferous forests, streamsides, on rocks and tree trunks; 2400–3600 m. Xizang, Yunnan.

Cystopteris tibetica is similar to C. moupperensis, but the lamina has short glandular hairs on both surfaces and the indusia also has short glandular hairs.

9. Cystopteris deqinensis Z. R. Wang, Acta Phytotax. Sin. 32(1): 84. 1994.

德钦冷蕨  de qin leng jue

Rhizomes long creeping. Fronds distant; fertile fronds 45–60 cm; stipe dark brownish at base, upward stramineous, 20–25 cm, 1.5–3 mm in diam., base clothed with brown hairs and few broadly ovate or lanceolate scales; lamina 3-pinnate, narrowly ovate or ovate-oblong, 25–35 Χ 15–20 cm, thinly herbaceous when dried, apex acuminate; pinnae up to 10–13 pairs or more, alternate, basal and second basal pinnae largest, subopposite, 5–7 cm apart from second basal pinnae, ascending, ovate-lanceolate, 12–17 Χ 4–6 cm, base slightly narrowed and inequilateral, with stalk 0.5–1.5 cm, apex acuminate; pinnules 10–15 pairs, anadromous, usually basiscopic one slightly longer than acroscopic one, basiscopic second and third basal pinnules largest, ovate to broadly lanceolate, 2.5–4.3 Χ 1.5–2.2 cm, base inequilateral, acroscopic base parallel to costae, basiscopic base broadly cuneate, with stalk 1.5–2 mm, apex acuminate, alternate; secondary pinnules 5–7 pairs, basal acroscopic pinnule largest, broadly ovate, 8–14 Χ 5–8 mm, base narrowly cuneate, pinnatifid, toothed at margin, apex rounded-obtuse; veins visible on both surfaces, forked, terminating in emarginations. Sori orbicular, 1–6 sori per ultimate segment, abaxial on veins; indusia pale green to brownish, orbicular or shallowly cupular, without glands, covering sporangia when young, hidden at maturity. Spores bean-shaped, perispore very densely echinate.

* On rocks in valleys, mountain slopes in Quercus forests, streamsides; 2400–3000 m. NW Yunnan.

10. Cystopteris moupperensis Franchet, Nouv. Arch. Mus. Hist. Nat., sιr. 2, 10: 111. 1887.

宝兴冷蕨  bao xing leng jue

Cystopteris mairei Brause; C. sphaerocarpa Hayata; C. sudetica A. Braun & Milde var. moupperensis (Franchet) C. Christensen; C. tangutica Grubov.

Rhizomes long creeping, 1–2 mm in diam., rhizomes and stipe bases clothed with brown hairs and few, pale brown, broadly ovate, membranous scales. Fronds distant; fertile fronds (20–)30–40(–50) cm; stipe stramineous or chestnut-brown, shiny, 10–22(–25) cm, 1–2 mm in diam.; lamina pinnate-pinnatifid to 3-pinnate, ovate or deltoid-ovate, 9–17(–25) Χ 5–8(–15) cm, thinly herbaceous when dried, apex acuminate; pinnae 8–12(–15) pairs, ascending, basal pinnae oblong or ovate-lanceolate, 5–10 Χ 2–3.5 cm, as long as and wider than second basal pinnae, toothed, base slightly narrowed and inequilateral, with stalk 3–4 mm (2–3 mm in second basal pinnae), apex acuminate, subopposite, 3–3.5(–5) cm apart from second basal pinnae; pinnules 8–12 pairs, anadromous, usually acroscopic pinnule slightly shorter than basiscopic one, second basal basiscopic pinnule largest, deltoid-ovate or narrowly triangular, 1–2.2(–3) cm Χ 8–13 mm, toothed, inequilateral, acroscopic base parallel to costae, basiscopic base broadly cuneate, with stalk 1–2 mm, apex obtuse, alternate; secondary pinnules or pinnule segments 3 or 4 pairs; basal acroscopic one largest, broadly ovate, 4–9 Χ 3–5 mm, base cuneate and adnate to costular wing, cut or toothed at margin, apex rounded-obtuse and toothed; upper secondary pinnules rhomboid or shortly linear, apex and acroscopic side toothed. Veins visible on surfaces, one to several times forked, terminating in emarginations. Sori small, orbicular, abaxial on suprabasal veins, 1 or 2 sori per ultimate segment; indusia pale green or yellow-brown, orbicular or shallowly cupular, membranous, without tiny glandular hairs, covering sporangia when young, hidden at maturity. Perispore echinate.

Damp areas and on wet rocks in mixed forests; 1000–4100 m. Gansu, NW Guizhou, Hebei, Henan, Qinghai, Shaanxi, Sichuan, Taiwan, SE Xizang, Yunnan [N India].

Cystopteris moupperensis is very similar to C. sudetica, but different in the indusia without tiny glandular hairs and the plants moderately stout. The two are segregated phytogeographically: C. sudetica occurs in NE and N China, and occasionally in high mountains of Yunnan and Xizang, while C. moupperensis occurs in mountains of W China, north to Qinling Shan and west to E Xizang.

11. Cystopteris pellucida (Franchet) Ching ex[??in] C. Christensen, Index Filic., Suppl. 3: 67. 1934.

膜叶冷蕨  mo ye leng jue

Aspidium pellucidum Franchet, Nouv. Arch. Mus. Hist. Hat., sιr. 2, 10: 119. 1887; Cystopteris alata Ching; Dryopteris pellucida (Franchet) C. Christensen; Nephrodium pellucidum (Franchet) Diels.

Rhizomes long creeping, ca. 2 mm in diam., rhizome and stipe bases densely clothed with red-brown short hairs, and few pale brown broadly ovate membranous scales. Fronds distant; fertile fronds (20–)50–60 cm; stipe pale stramineous or brownish red, shiny, (10–)20–32 cm, 1–2 mm in diam., with few pale brown, broadly ovate, membranous scales mixed with short hairs at base; lamina 3-pinnate, ovate to narrowly ovate-oblong, (10–)20–33 Χ (5–)10–15(–25) cm, thinly herbaceous or submembranous when dried, apex long acuminate; pinnae (10–)12–15(–17) pairs, ascending, basal pinnae longest, deltoid-lanceolate, (5–)8–14 Χ 2.5–4.5 cm, base inequilateral, with stalk 3–5(–8) mm, apex caudate-acuminate, subopposite, (3–)5(–7) cm apart from second basal pinnae; pinnules 8–12 pairs, anadromous, alternate, usually acroscopic one slightly smaller than basiscopic one, second basal basiscopic pinnules largest, narrowly ovate or narrowly triangular, 1.5–2.5(–4) Χ 1–1.5 cm, base very inequilateral, acroscopic base narrowly cuneate, parallel to costae, abaxial base broadly cuneate, very shortly stalked or subsessile, apex acute or subobtuse and toothed; secondary pinnules or pinnule segments 3–5 pairs, basal acroscopic one largest, oblong or ovate, rounded-obtuse or subtruncate at apex, toothed at apex and margin; veins visible on surfaces, more obvious abaxially, simple or 1- or 2-forked, terminating in emarginations. Sori brown, orbicular, abaxial suprabasally, 1 or 2 sori per ultimate segment; indusia pale brown, orbicular or half cup-shaped, membranous, without tiny glandular hairs, covering sporangia when young, hidden at maturity. Perispore echinate. 2n = ca. 84.

* Forest understories on mountain slopes, damp areas at streamsides; 1500–3700 m. S Gansu, Henan, Shaanxi, W Sichuan, Xizang, Yunnan.

Cystopteris pellucida is very similar to and sympatric with C. moupperensis, but differs from it in the plant is slightly larger, the ultimate segments wider, and lamina thinner. The indusia are larger and attached up to 2/3 of the circumference of the receptacle.



[1] Department of Botany, National Museum of Nature and Science, Tsukuba 305-0005, Japan.

[2] Department of Botany, Duke University, Box 90338, Durham, North Carolina 27708-0338, U.S.A.