WOODSIACEAE [Draft]

岩蕨科  yan jue ke

Zhang Gangmin (张钢民)[1]; Masahiro Kato[2], Alexander Shmakov[3]

Lithophytes or xerophytes, small or medium-sized. Rhizomes short, erect, decumbent, or ascending, simple dictyostelic, covered with scales. Fronds clustered, monomorphic, deciduous or sometimes evergreen. Stipe mostly shorter than lamina, ± scaly and hairy, articulate or not articulate. Lamina 1-pinnate to bipinnatifid, elliptic-lanceolate to narrowly lanceolate, herbaceous or papery, frequently covered with articulate multicellular hairs, sometimes with glandular hairs or capitate glands. Veins free, pinnate, usually ending in enlarged hydathodes. Sori round, consisting of 3–18 sporangia, borne at middle or apex of veinlets. Indusia inferior, saucer-shaped to cup-shaped, margin long ciliate, or indusia absent, degenerated into curly, multicellular hairs (Woodsia), or sphaeropteroid (bladdery) with openings at tip (Protowoodsia), sometimes also covered with false indusia, i.e., reflexed leaf margins (Cheilanthopsis). Spores ellipsoidal or somewhat spherical, monolete, wall surface folded, cristate, tuberculate, or echinate.

About four genera (including the Caucasian Hymenocystis), ca. 43 species; widely distributed in the north temperate and frigid zones, rarely in Central and South America, Africa (Angola, South Africa) and Madagascar; three genera and 23 species in China, mainly in northern regions (eight endemic).

The current classification of the genera requires evaluation by molecular phylogenetic evidence.

1a.       Stipe obviously articulate; indusia saucer- to cup-shaped, margin long ciliate
                                                                                                                                                             3. Woodsia

1b.       Stipe continuous; indusia spherical or degenerated into curly, multicelluar hairs.

2a.       Indusia platelike with unequal lobes or dissected with filiform lobes .....................  3. Woodsia

2b.       Indusia spherical, tearing when mature.

3a.       Lamina glabrous or with few glandular trichomes, margin flat, not reflexed; pinnae not articulate to rachis                                                                                                                  1. Protowoodsia

3b.       Lamina pubescent, margin reflexed to form false indusia (usually fallen later); pinnae articulate to rachis                                                                                                                 2. Cheilanthopsis

1. PROTOWOODSIA Ching, Lingnan Sci. J. 21: 36. 1945.

膀胱蕨属  pang guang jue shu

Lithophytes, small. Rhizomes erect, short, apex densely scaly; scales brown, lustrous, ovate-lanceolate or lanceolate, entire. Fronds clustered or subclustered, usually diverging; stipe short, not articulate, brittle, usually lower parts persistent. Lamina 1-pinnate-pinnatifid, lanceolate, much longer than stipe, herbaceous, glabrous or glandular hairs sparse. Veins free, pinnate, not reaching laminar margin. Sori small, round, consisting of 6–12 sporangia, medial or submarginal. Indusia inferior, spherical or bladdery, completely wrapping sporangia, tearing apically at maturity. Spores ellipsoidal, monolete, perispores corrugate, exine glabrous on surface. Χ = 33.

One species in Asia; the one species in China.

1. Protowoodsia manchuriensis (Hooker) Ching, Lingnan Sci. J. 21: 36[??37]. 1945.

膀胱蕨  pang guang jue

Woodsia manchuriensis Hooker, Second Cent. Ferns t. 98. 1861; Diacalpe manchuriensis (Hooker) Trevisan; Physematium manchuriensis (Hooker) Nakai.

Plants (8–)15–20 cm. Rhizomes short, erect; scales brown, lustrous, ovate-lanceolate or lanceolate, entire. Fronds clustered; stipe brownish stramineous, 2–2.5 cm, less than 1 mm in diam., brittle, usually breaking, sparsely short glandular hairy, with few scales in lower part. Lamina 1-pinnate-pinnatifid, lanceolate, 12–18 Χ 1.5–4 cm, herbaceous, sparsely glandular hairy, base attenuate, apex acuminate. Pinnae (12–)16–20 pairs, oblique, subsessile; middle pinnae largest, ovate-lanceolate, 1–1.5 Χ up to 1 cm, pinnatifid, apex obtuse; lower pinnae shorter than above, widely separate; basal pair of pinnae ovate or flabellate. Ultimate segments oblong, undulate or dentate. Veins free, pinnate, not reaching laminar margin. Sori consisting of 6–8 sporangia, medial or submarginal. Indusia prominent, spherical, thinly membranous. 2n = 66.

On rocks in forests; 200–4000 m. Anhui, Guizhou, Hebei, Heilongjiang, Henan, Jiangxi, Jilin, Liaoning, Sichuan, Shandong, Zhejiang [Japan, Korea, Russia].

2. CHEILANTHOPSIS Hieronymus, Notizbl. Bot. Gart. Mus. Berlin-Dahlem 7: 406[??409]. 1920.

滇蕨属  dian jue shu

Lithophytes, middle-sized or small. Rhizomes ascending or decumbent, apex covered with brown, membranous lanceolate scales. Fronds clustered; stipe short, brittle, usually broken, base or lower portion scaly, upper portion pubescent. Lamina 1-pinnate-pinnatifid, lanceolate, gradually tapering to both ends, herbaceous, covered with multicelluar glandular hairs. Pinnae lanceolate, sessile, articulate to rachis. Ultimate segments hemielliptic. Veins free, pinnate, not reaching laminar margin. Sori small, consisting of 4–10 sporangia, submarginal, at veinlet endings, round. Indusia inferior, subspherical, sometimes not fully developed, thinly membranous; false indusia formed by pale thin, reflexed laminar margins. Spores ellipsoidal, monolete, wall-surface folded, granulate. Χ = 37, 41.

Three species in Himalaya and China; the three species in SW China (one endemic).

1a.       Articulate hairs dense on both surfaces of lamina; indusia not large, false indusia conspicuous, round-hemiovate, nearly reaching midvein .................................................................. 1. C. indusiosa

1b.       Articulate hairs sparse on both surfaces of lamina; indusia large, globose, false indusia small, reniform or cordate, far from midvein.

2a.       Stipe densely scaly at base, upward with dense articulate hairs, rachis also articulate-hairy  2. C. elongata

2b.       Stipe sparsely scaly, rachis with dense glandular hairs ............................  3. C. kangdingensis

1. Cheilanthopsis indusiosa (H. Christ) Ching, Sinensia 3(5): 154. 1932.

滇蕨  dian jue

Woodsia indusiosa H. Christ in Lecomte, Notul. Syst. (Paris) 1: 44. 1909; Cheilanthes straminea Brause; Cheilanthopsis straminea (Brause) Hieronymus ex Copeland.

Plants 25–50 cm. Rhizomes ascending, apex densely covered with scales; scales brown, broadly lanceolate, up to 1 cm, entire. Fronds clustered; stipe stramineous, 3–11 cm, 1–2 mm in diam., brittle, densely covered with scales, articulate hairs sparse when young, deciduous, middle and upper parts glabrous. Lamina 1-pinnate-pinnatifid, lanceolate, 25–40 Χ 5–6.5 cm, herbaceous, densely covered with articulate hairs and glands on both sides, base attenuate, acuminate; rachis pale stramineous, with articulate hairs and sparse brown, narrowly lanceolate scales. Pinnae 25–30 pairs, spreading or slightly oblique, sessile; lower pinnae reduced to lobes, lobes triangular, ligulate or auriculate, remote; middle pinnae largest, lanceolate, 2–3.5 cm Χ 3–10 mm, base truncate, pinnatifid, apex obtuse; lobes 7–12 pairs, opposite, spreading, hemielliptic; basal lobes largest, 3–5 mm, undulate. Veins free, obscure. Sori consisting of 4–8 sporangia, located at vein endings, submarginal. Indusia not conspicuous, thinly membranous. False indusia hemiovate, large, membranous, nearly reaching midvein, fimbriate. 2n = 74.

On rocks in broad leaved forests; 2100–3200 m. SW Sichuan, NW Yunnan [Bhutan].

2. Cheilanthopsis elongata (Hooker) Copeland, Univ. Calif. Publ. Bot. 12: 395. 1931.

长叶滇蕨  chang ye dian jue

Woodsia elongata Hooker, Sp. Fil. 1: 62. 1844; Physematium elongata (Hooker) Trevisan.

Plants 25–28 cm. Rhizomes prostrate, apex densely covered with scales; scales brown, broadly lanceolate, 4–6 mm, membranous, entire. Fronds clustered; stipe stramineous, 3–6 cm, 1–2 mm in diam., brittle, base densely covered with scales, upward and rachises densely covered with articulate hairs, and with few brown lanceolate scales. Lamina 1-pinnate-pinnatifid, lanceolate, 18–22 cm Χ 2.5–4 cm, thinly herbaceous, both surfaces with sparse articulate hairs and glands, base attenuate, acuminate. Pinnae 17–21 pairs, remote, spreading; lower pinnae shortened to deltoid-ovate to auricular lobes; middle pinnae largest, broadly lanceolate, 1.2–2 cm Χ 6–10 mm, base truncate, pinnatifid, apex obtuse; lobes 5 or 6 pairs, opposite, spreading, hemielliptic; basal lobes slightly larger, 3–5 Χ 1.5–3 mm, margin undulate. Sori consisting of 4–6 sporangia, terminal at vein tips, submarginal. Indusia spherical, thinly membranous. False indusia broadly reniform or cordate, small, membranous, remote from each other and from midvein. 2n = 82.

Rock crevices in forests; 3200–3400 m. Xizang, NW Yunnan [Bhutan, Myanmar, India, Nepal].

3. Cheilanthopsis kangdingensis (H. S. Kung, L. B. Zhang & X. S. Guo) Shmakov in Chandra & Srivastava (eds.), Pterid. New Millen. 64. 2003.

康定岩蕨  kang ding yan jue

Woodsia kangdingensis H. S. Kung, L. B. Zhang & X. S. Guo, Acta Bot. Yunnan. 17: 421. 1995.

Plants 16–23 cm. Rhizome erect, apex scaly; scales brown, ovate-lanceolate, 4–6 mm, margin entire. Fronds clustered; stipe stramineous, 2–4 cm, glabrous, sparsely scaly; scales brown, lanceolate or narrowly ovate, entire. Lamina 2-pinnatifid, narrowly lanceolate, 12–20 Χ 1.8–3 cm, base attenuate, apex acuminate; rachis with dense, short glandular hairs. Pinnae 14–19 pairs, subopposite, triangular-ovate, 8–18 Χ 4–8 mm, obtuse, base subequal, pinnatifid. Pinna-lobes 4–7 pairs, herbaceous, obtusely dentate; veins inconspicuous. Sori round; indusia membranous, apex erose.

* Rocks and rock crevices in forests; 3400–3800 m. Sichuan.

3. WOODSIA R. Brown, Prodr. 158. 1810, [‘Woodia’].

岩蕨属  yan jue shu

Plants small, usually on rocks. Rhizomes short, erect or ascending, rarely decumbent, scaly; scales lanceolate or linear-lanceolate, membranous. Fronds clustered; stipe shorter than lamina, articulate at various leves, i.e., from top to near base, or continuous in some species. Lamina 1-pinnate or 1-pinnate-pinnatifid, lanceolate, herbaceous or subpapery, usually with articulate hairs or scales, rarely glabrous, base gradually tapering. Veins free, pinnate, not reaching laminar margin. Sori small, consisting of 3–18 sporangia, round, on veinlets or at veinlet tips. Indusia inferior, various, usually saucer- to cup-shaped, dissected into ciliate platelike segments or reduced into curly, multicelluar hairs, sometimes exindusiate. Spores ellipsoidal, monolete, perispore folded with granulate, echinate or tuberculate ornamentations on surface. Χ = 39, 41.

About 38 species: widely distributed in north temperate regions and higher elevations of tropical America, Africa, and Madagascar; 19 species in China (seven endemic).

1a.       Stipe continuous, i.e., not articulate; indusia reduced into curly, multicelluar hairs or apparently exindusiate (subg. Eriosorus).

2a.       Rhizome ascending or decumbent; lamina 1-imparipinnate.

3a.       Pinnae entire or slightly undulate; sori exindusiate or with much reduced indusia hidden  18. W. cycloloba

3b.       Pinnae lobed to pinnatifid; indusia consisting of pale, long, crested hairs
                                                                                                                 19. W. okamotoi

2b.       Rhizome erect, occasionally ascending; lamina 1- paripinnate or bipinnatifid, apical pinna absent, pinnae not entire; sori covered by curly, long multicellular hairs.

4a.       Lamina 1-pinnate.

5a.       Rachis with sparse long hairs, occasionally with one or two scales; pinnae ovate, both surfaces sparsely shortly hairy ........................................................................  13. W. macrospora

5b.       Rachis densely covered with scales; pinnae oblong-ovate, both surfaces densely covered with loose long hairs ........................................................................................  14. W. cinnamomea

4b.       Lamina bipinnatifid.

6a.       Plant 2–10 cm tall (usually ca. 5 cm tall); stipe and rachis of black wood color; laminae covered with hairs and scales abaxially; pinnae 4-10 pairs
.................................................................................................................  15. W. lanosa

6b.       Plant to 25 cm; lamina densely covered only with loose ferruginous hairs; stipes brown or brown-stramineous; pinnae 6--15 pairs.

7a.       Pinnae ovate, basal pinnae not reduced or slightly reduced; indusia composed of glandular-tipped hairs .................................................................................................  16. W. andersonii

7b.       Pinnae elliptic, lower pinnae obviously reduced; indusia composed of ciliate platelike lobes             17. W. rosthorniana

1b.       Stipe articulate at various levels; indusia cup- or saucer-shaped, fimbriate, long ciliate.

8a.       Stipe articulate at or below middle (subg. Woodsia).

9a.       Lamina covered with long hairs and scales; stipes robust, castaneous or castaneous-brown.

10a.     Lamina subglabrous, with sparse long hairs and scales on rachis; pinna short, triangular-ovate or ovate, pinnatifid .....................................................................................................  4. W. alpina

10b.     Lamina sparsely long hairy, rachis and costae scaly beneath; pinna long, ovate-lanceolate, pinnatipartite nearly to costa ...........................................................................................  5. W. ilvensis

9b.       Lamina glabrous or with long hairs, without any scales; stipes slender, substraminous.

11a.     Lamina sparsely hairy abaxially, hairs long; pinna shortly stalked
............................................................................................................  3. W. shensiensis

11b.     Lamina glabrous; pinna sessile.

12a.     Pinna triangular-ovate, apex obtuse, 3–5 mm ..........................................  1. W. glabella

12b.     Pinna nearly rhombic, apex acuminate or acute, up to 8 mm ..................  2. W. hancockii

8b.       Stipe articulate at top, or sometimes at upper parts.

13a.     Indusia platelike; base of pinna usually symmetrical or subsymmetrical (subg. Woodsia).

14a.     Stipe 4–8 cm, brown or light castaneous; pinna 1–2 cm, pinnatifid ......  11. W. subcordata

14b.     Stipe 1–3 cm, stramineous; pinna 4–6 mm, pinnatifid into 3–5 segments ......  12. W. sinica

13b.     Indusia cup-shaped; base of pinna asymmetrical or rarely symmetrical (subg. Acrolysis).

15a.     Pinnae below laminar apex free, elliptic-lanceolate or falcate-lanceolate; midvein scaly beneath.

16a.     Pinna entire or undulate .................................................................  6. W. polystichoides

16b.     Pinna pinnatilobate on both sides ................................................................  7. W. pilosa

15b.     Upper pinnae adnate, triangular-lanceolate or elliptic; midvein without scales or with few scales only beneath.

17a.     Only basal pinnae free, upper pinnae adnate, base symmetrical; rachis densely covered with long hairs, but without scales ................................................................  8. W. macrochlaena

17b.     Lower pinnae free, upper pinnae adnate, base asymmetrical; rachis covered with hairs and scales.

18a.     Pinna sparsely hairy, herbaceous ..........................................................  9. W. oblonga

18b.     Pinna densely hairy on both surfaces, subpapery .............................  10. W. intermedia

1. Woodsia glabella R. Brown ex Richardson in Franklin, Narr. Journey Polar Sea, 754. 1823.

光岩蕨  guang yan jue

Woodsia alpina (Bolton) Gray var. glabella (R. Brown ex Richardson) D. C. Eaton; W. hyperborea (Liljeblad) R. Brown var. glabella (R. Brown ex Richardson) Watt; W. lapponica Angstrom; W. yazawai Makino.

Plants 5–10 cm. Rhizomes compact, ascending, scaly as stipe base; scales concolorous, brown, lanceolate or ovate lanceolate, 2–3 mm, membranous, entire. Fronds clustered; stipe brown-stramineous, slender, 1–2 cm, brittle, articulate below middle, glabrous or rarely with few brown, linear scales on apical portion. Lamina 1-pinnate-pinnatifid, linear-lanceolate, 3–6 cm Χ 7–11 mm, thinly herbaceous, both surfaces glabrous or with occasional hairs, base attenuate, apex acuminate; rachis glabrous. Pinnae 4–9 pairs, well-separated, sessile, spreading; lower pinnae shorter than above, basal one flabellate; medial pinnae largest, deltoid-ovate, 3–5 Χ 2–5 mm, base truncate, pinnatifid, with 1–3 pairs of pinnules, apex obtuse. Pinnules elliptic or tongue-shaped, proximal pairs largest, ca. 3 mm, undulate or apically crenulate. Veins pinnate, visible. Sori consisting of few sporangia, submarginal. Indusia saucer-form, fimbriate, thinly membranous, brittle, usually fallen when mature. 2n = 78.

In rock crevices in needle-leaf forests and mixed forests; 2100–3700 m. Gansu, Hebei, Jilin, Qinghai, Xinjiang, Yunnan [Japan, Russia; Europe, North America].

2. Woodsia hancockii Baker, Ann. Bot. 5: 196. 1891.

华北岩蕨  hua bei yan jue

Woodsia gracillima C. Christensen; W. tsurugisanensis Makino.

Plants 3–10 cm. Rhizomes short, erect, scaly; scales concolorous, brown, ovate-lanceolate or elliptic, membranous. Fronds clustered; stipe stramineous, 1–2 cm, brittle, articulate below middle. Lamina 1-pinnate-pinnatifid, lanceolate, 2–8 cm Χ 3–12 mm, thinly herbaceous, both surfaces glabrous, base attenuate, apex acuminate. Pinnae 7–14 pairs, sessile, spreading or slightly oblique; lower pinnae shorter than above, remote from each other; middle pinnae largest, subrhombic or obliquely ovate, 4–8 Χ 3–5 mm, base broadly cuneate or truncate, pinnatifid, with 2 or 3 pairs of segments, apex acuminate or acute. Pinna segments obovate or tongue-shaped; basal pair largest, ca. 2 mm, undulate or with 1 or 2 crenations at tip. Veins pinnate, not reaching laminar margin. Sori round, consisting of a few sporangia, located at veinlet tips or on veinlets, usually 1–3 per segment. Indusia saucer-form, thinly membranous, margin with flexuose brown articulate long hairs.

Wet shaded rock crevices in forests; 1700–2200 m. Hebei, Heilongjiang, Jilin, Nei Mongol, Shaanxi, Shanxi [Japan, Korea, Russia].

Woodsia hancockii is similar to W. glabella in the slender and glabrescent fronds, but differs from it in the more slender leaf, trapeziform pinnae, and much larger indusia fringed with long hairs.

3. Woodsia shensiensis Ching, Sinensia 3(5): 141. 1932.

陕西岩蕨  shan xi yan jue

Plants 10–15 cm. Rhizomes short, erect, scaly; scales light brown, lanceolate, ca. 3 mm, membranous, entire. Fronds clustered; stipe stramineous, 3–5 cm, articulate at lower part, scaly at base, upward sparsely scaly; scales narrowly lanceolate. Lamina 1-pinnate-pinnatifid to 2-pinnate, lanceolate, 7–10 Χ 1.5–2.9 cm, thinly herbaceous, abaxially with few brown articulate hairs, adaxially glabrous, base attenuate, apex acuminate; rachis, costae, and leaf margin also with few similar hairs. Pinnae 8–10 pairs, ca. 1 cm apart, shortly stalked, slightly oblique; lower pinnae shorter than above; middle pinnae largest, ovate-rhombic, 1–1.5 cm Χ 7–10 mm, base broadly cuneate, pinnatifid or pinnate, with 2 or 3 pairs of pinnules, apex acute. Pinnules obovate, basal pair largest, ca. 7 mm, serrate or crenate. Veins pinnate, distinct. Sori subround, usually terminal at ends of veinlets, 2–4 per segment. Indusia saucer-form, with long-villous margin.

* On rocks in forests; 2000–2900 m. Nei Mongol, Shaanxi, E Sichuan.

4. Woodsia alpina (Bolton) Gray, Nat. Arr. Brit. Pl. 2: 17. 1821.

西疆岩蕨  xi jiang yan jue

Acrostichum alpinum Bolton, Fil. Brit. 76. 1790; A. hyperboreum Liljeblad; Polypodium arvonicum Withering; Woodsia himalaica Ching & S. K. Wu; W. hyperborea (Liljeblad) R. Brown; W. ilvensis (Linnaeus) R. Brown var. alpina (Bolton) Watt.

Plants 8–12 cm. Rhizomes compact, ascending, with cluster of persistent stipe bases of ± equal length; scales concolorous, brown, lanceolate, ca. 5 mm, membranous, fimbriate. Fronds clustered; stipe reddish brown, lustrous, 1–2 cm, ca. 1 mm in diam., articulate below middle, sparsely scaly, and scales later fallen. Lamina 1-pinnate-pinnatifid, lanceolate, 5.5–9 Χ 1–1.7 cm, herbaceous or thinly herbaceous, both surfaces glabrous, base attenuate, apex acuminate; rachis with sparse long hairs and linear-lanceolate scales. Pinnae 9–11 pairs, separate, oblique; lower pinnae shorter than above, ovate, 2–3 mm; middle pinnae largest, ovate or deltoid-ovate, 5–9 Χ 4–7 mm, pinnatifid, with 3 or 4 pairs of lobes, acute. Pinna-lobes hemielliptic, 2–3 mm, margin usually involute when dried. Veins pinnate, ending in hydathodes close to laminar margin, obvious adaxially. Sori round. Indusia saucer-form, with fimbriate long-ciliate margin. 2n = 156.

Rock crevices in forests; ca. 4000 m. Xizang (H. C. Levinge’s collection) [Russia; Europe, North America].

H. C. Levinge’s collection is the only record of Woodsia alpina from NW Himalaya. It is closely comparable to northern European forms. It has always been identified as W. subcordata or other allied species, and cited by some authors in the flora of N and NE China, and the vicinity.

5. Woodsia ilvensis (Linnaeus) R. Brown, Prodr. 158. 1810.

岩蕨  yan jue

Acrostichum ilvense Linnaeus, Sp. Pl. 2: 1071. 1753; Nephrodium rufidulum Michaux; Polypodium ilvense (Linnaeus) Swartz [??(Linnaeus) Villars].

Plants 12–17 cm. Rhizomes erect or ascending, scaly; scales concolorous, brown, broadly lanceolate, ca. 4 mm, membranous, entire. Fronds clustered; stipe castaneous, lustrous, 3–7 cm, ca. 1 mm in diam., with long hairs and linear-lanceolate scales, articulate below middle. Lamina 1-pinnate-pinnatifid, lanceolate, 8–11 Χ 1.3–2 cm, herbaceous, covered with articulate hairs, especially abaxially, base gradually attenuate, apex acuminate; rachis and costae with brown, linear scales and long articulate hairs. Pinnae 10–20 pairs, sessile, well-separated, oblique; middle pinnae largest, ovate-lanceolate, 8–11 Χ 4–8 mm, pinnatifid, with 3–5 pairs of lobes, acute. Pinna-lobes elliptic, basal pair largest, 2–4 mm, entire or irregularly undulate. Veins pinnate, veinlets not reaching laminar margin. Sori rounded and attached at ends of veinlets, submarginal. Indusia saucer-form, membranous, margin long ciliate. 2n=78(82).

Cliffs and rocky slopes; 200–2200 m. Hebei, Heilongjiang, Jilin, Nei Mongol, Xinjiang [Kazakhstan, Mongolia, Russia; Europe, North America].

6. Woodsia polystichoides D. C. Eaton, Proc. Amer. Acad. Arts 4: 110. 1858 [??1860].

耳羽岩蕨  er yu yan jue

Woodsia brandtii Franchet & Savatier; W. polystichoides var. incisa Ching & Y. T. Hsien; W. polystichoides var. nudiuscula Hooker; W. polystichoides var. veitchii Hance.

Plants 15–30 cm. Rhizomes short, erect, scaly; scales brown, lanceolate or ovate-lanceolate, ca. 4 mm, membranous, entire. Fronds clustered; stipe stramineous or brown-stramineous, slightly lustrous, 4–12 cm, 1–1.5 mm in diam., articulate at or near top, with linear or linear-lanceolate scales and long multicelluar hairs. Lamina linear-lanceolate or narrowly lanceolate, 10–23 Χ 1.5–3 cm, papery or herbaceous, abaxially with mixture of linear scales and long hairs, adaxially glabrous or with few long hairs, base attenuate, 1-pinnate, apex acuminate; rachis long hairy. Pinnae 16–30 pairs, sessile, spreading or slightly oblique, entire or undulate; lower 3 or 4 pairs of pinnae reduced and reflexed downward, basal pair of pinnae triangular; middle pinnae largest, elliptic-lanceolate, slightly falcate, 8–20 Χ 4–7 mm, base asymmetric, acroscopically auriculate, truncate, basicopically cuneate, apex acute. Veins obvious, pinnate, veinlets 2-forked, ending in submarginal hydathodes. Sori round, terminal at tips of acroscopic veinlets, submarginal. Indusia cup-shaped, shallowly lobed, ciliate. 2n = 82.

Crevices of shaded rocks; 200–2700 m. Widely distributed in C, E (including Taiwan, but not in Fujian), N, NW, SW (Sichuan, Yunnan) China [Japan, Korea, Russia].

7. Woodsia pilosa Ching, Fl. Tsinling. 2: 222. 1974.

嵩县岩蕨  song xian yan jue

Plants ca. 20 cm tall. Rhizomes erect, scaly at apex; scales light brown, lanceolate, 4–5 mm, thinly membranous, fimbriate. Fronds clustered; stipe light stramineous, lustrous, 5–7 cm, ca. 1.2 mm in diam., articulate at top, base densely scaly, upward glabrous. Lamina 1-pinnate-pinnatifid, linear-lanceolate, 11–12 Χ 1.6–1.8 cm, herbaceous, both surfaces densely covered with articulate hairs and occasionally abaxially with elongate scales along midvein, apex acuminate; rachis with mixture of linear scales and multicellular hairs. Pinnae 15–18 pairs, free, sessile, spreading; lower pinnae opposite, slightly shortened; middle pinnae largest, ovate-lanceolate, 7–10 Χ 3–4 mm, alternate, shallowly pinnatifid, base asymmetric, ± auriculate, acroscopically truncate, basiscopically cuneate, apex obtuse. Ultimate segments elliptic, touching each other, entire. Veins pinnate, invisible. Sori round, terminal at tips of acroscopic veinlets, submarginal. Indusia cup-shaped, margin ciliate, shallowly lobed into 3 segments at maturity.

* Rock crevices. Henan (Song Xian).

8. Woodsia macrochlaena Mettenius ex Kuhn, J. Bot. 6: 270. 1868.

大囊岩蕨  da nang yan jue

Woodsia frondosa H. Christ; W. japonica Makino.

Plants 5–16(–20) cm. Rhizomes short, erect or ascending, scaly at apex; scales brown, 3–4 mm, lanceolate, membranous. Fronds clustered; stipe stramineous, 1–7.5 cm, ca. 1 mm in diam., articulate at top, with sparse, brown, articulate hairs. Lamina 2-pinnatifid, elliptic-lanceolate, 4–10 Χ 2–3 cm, herbaceous, densely covered with articulate hairs, especially abaxially, base slightly narrowed or not, apex acute; rachis similarly hairy. Pinnae 7–10 pairs, connected to each other by wings except for basal pinnae, opposite, spreading or slightly oblique, sessile; middle pinnae slightly longer, elliptic-oblong, 7–11(–22) Χ 5–1.2 mm, base symmetric, pinnatifid, lobes entire, apex obtuse. Veins obscure, pinnate, tips often enlarged to form hydathodes. Sori round, terminal at tips of veinlets, submarginal. Indusia cup-shaped, membranous, lacerate.

Rock crevices in forests. Hebei, Liaoning, Shandong [Japan, Korea, Russia].

9. Woodsia oblonga Ching & S. H. Wu, Fl. Tsinling. 2: 221. 1974.

妙峰岩蕨  miao feng yan jue

Plants 7–18 cm. Rhizomes ascending, scaly at top; scales light brown, ca. 3 mm, lanceolate, thinly membranous, margin ciliate. Fronds clustered; stipe brown-stramineous, lustrous, 2–5 cm, ca. 1 mm in diam., articulate at top, with sparse, curly, long hairs and linear scales. Lamina 1-pinnate, lanceolate, 6–16 Χ 2–3.5 cm, herbaceous, both surfaces with sparse, brown, articulate hairs, base slightly attenuate, apex acute; rachis stramineous, with mixture of linear scales and articulate hairs. Pinnae 8–18 pairs, sessile, spreading; lower 1 or 2 pairs of pinnae slightly shortened and reflexed downward; middle pinnae larger, elliptic, 1–1.5 cm Χ 5–8 mm, base asymmetric, acroscopically truncate, close to rachis, margin nearly entire or slightly undulate, apex obtuse; upper pinnae connected to each other by wings. Veins pinnate, not reaching laminar margin. Sori round, terminal at ends of veinlets, nearly marginal. Indusia cup-shaped, margin ciliate, shallowly lobed into 2 or 3 segments at maturity.

* Crevices of shaded rocks; 200–1800 m. Hebei, Henan, Shandong.

10. Woodsia intermedia Tagawa, Acta Phytotax. Geobot. 5: 250. 1936.

东亚岩蕨  dong ya yan jue

Woodsia subintermedia Tzvelev, [nom. nov. replaces W. intermedia Tagawa]; W. taishanensis F. Z. Li & C. K. Ni.

Plants 10–25 cm. Rhizomes short, erect or ascending, scaly; scales brown, lanceolate, ca. 3 mm, membranous, margin subentire or sparsely ciliate. Fronds clustered; stipe brown-stramineous or light chestnut, 3–7.5 cm, ca. 1 mm in diam., articulate at top, with curly, long hairs and brown, linear scales later deciduous. Lamina 1-pinnate, lanceolate, 8–18 Χ 2–3.8 cm, papery, both surfaces densely hairy, base slightly attenuate, apex acuminate or acute; rachis with mixture of linear scales and articulate hairs. Pinnae 14–20 pairs, sessile, spreading, well-separated from each other; lower pairs of pinnae shorter than above, hemielliptic or deltoid-ovate; middle pinnae largest, deltoid-lanceolate, 1–2 cm Χ 4–10 mm, base broadly cuneate, asymmetric, acroscopically obviously auriculate, undulate or crenate, apex obtuse; upper pinnae connected to each other by wings. Veins obscure, veinlets oblique, 2- or 3-forked, ending with hydathodes and not reaching laminar margin. Sori round, terminal at vein tips, submarginal. Indusia cup-shaped, margin ciliate or hairy.

Crevices of shaded rocks in forests; 500–1800 m. Hebei, Heilongjiang, Henan, Jilin, Liaoning, Shandong, Shanxi [Japan, Korea, Russia].

11. Woodsia subcordata Turczaninow, Bull. Soc. Imp. Naturalistes Moscou 5: 206. 1823 [??1832].

等基岩蕨  deng ji yan jue

Woodsia conmixta Ching; W. eriosora H. Christ; W. kitadakensis Ohwi; W. longifolia Tagawa; W. pilosella Ruprecht; W. polystichoides D. C. Eaton f. sinuata (Hooker) Kitagawa; W. polystichoides var. sinuata Hooker; W. viridis Ching.

Plants 14–22 cm. Rhizomes short, erect or ascending, scaly; scales brown, ovate-lanceolate or broadly lanceolate, ca. 4 mm, membranous, margin ciliate, apex acuminate. Fronds clustered; stipe brownish or brown-stramineous, polished, (2–)4–8 cm, ca. 1 mm in diam., with oblique or horizontal articulation at top, base scaly, upper parts with sparse articulate long hairs and linear scales, later deciduous. Lamina 1-pinnate-pinnatifid, lanceolate, 8–15 Χ 2–3 cm, papery, both surfaces sparsely hairy and scaly, or occasionally subglabrous, hairs gray or grayish brown, long-articulate; scales brown, linear, small; lamina base slightly attenuate, apex obtuse or acuminate; rachis stramineous or tinted castaneous-brown, as hairy and scaly as stipe. Pinnae 11–16 pairs, subopposite or alternate, lower 2 or 3 pairs of pinnae gradually shortened, basal pair usually ca. 5 mm; middle pinnae largest, hemielliptic or narrowly triangular-lanceolate, 8–20 Χ 4–10 mm, base rounded-truncate, acroscopically auriculate, auricles ± overlapping rachis, pinnatifid to 1/2, apex obtuse; segments 3 or 4 pairs, hemielliptic, 2–4 mm, entire, rounded. Veins faintly visible, simply pinnate in segments, veinlets oblique, ending with hydathodes. Sori round, terminal at veinlet tips, submarginal. Indusia saucer-shaped, margin ciliate.

In rock crevices in forests; 500–3000 m. Hebei, Heilongjiang, Jilin, Liaoning, Nei Mongol, Shanxi [Japan, Korea, Mongolia, Russia].

12. Woodsia sinica Ching, Sinensia 3: 145. 1932.

山西岩蕨  shan xi yan jue

Plants 8–9 cm. Rhizomes short, erect, scaly; scales brown, ovate-lanceolate, ca. 3 mm, membranous, margin ciliate. Fronds clustered; stipe stramineous, 1–3 cm, ca. 1 mm in diam., articulate at top or above middle, with sparse articulate hairs and few lanceolate scales, later fallen. Lamina 1-pinnate-pinnatifid, linear-lanceolate, 4.5–6 cm Χ 8–12 mm, thinly papery, abaxially with few articulate hairs, adaxially glabrous, base slightly attenuate, apex acuminate; rachis sometimes with sparse linear scales. Pinnae 12–14 pairs, sessile, well-separated from each other; lower 2 pairs of pinnae reduced; middle pinnae equally long, triangular-ovate, 4–6 Χ 3–4 mm, base rounded-cuneate, pinnatifid, apex acute. Ultimate segments 3–5 pairs, basal one largest, hemielliptic, undulate, apex rounded. Veins obscure, almost reaching laminar margin. Sori round, terminal at end of veinlets, 3 or 4 per pinna; indusia saucer-shaped, fimbriate.

* Rock crevices. Shanxi (Ningwu).

13. Woodsia macrospora C. Christensen & Maxon, J. Wash. Acad. Sci. 17: 499. 1927.

甘南岩蕨  gan nan yan jue

Plants 12–17 cm. Rhizomes ascending, scaly; scales brown, lanceolate, ca. 3 mm, membranous. Fronds clustered; stipe dark stramineous, 6–9 cm, ca. 1 mm in diam., brittle, with sparse articulate hairs and few linear scales, upper part of stipes usually fallen, middle and lower parts persistent, clustered (articulation not seen). Lamina 1-pinnate, linear-lanceolate, herbaceous, pubescent, abaxial surface more densely pubescent, gradually attenuate to both ends; rachis with mixture of sparse articulate hairs and linear scales. Pinnae 10 or 11 pairs, sessile, well-separated, spreading; lower 1 or 2 pairs slightly shortened, ovate, ca. 4 mm; middle pinnae largest, suborbicular-ovate, 5–6 Χ 3–5 mm, base cuneate, margin irregularly undulate, slightly involute, apex obtuse. Veins obscure, pinnate, nearly reaching laminar margin. Sori round, medial, 3–5 per pinna. Indusia consisting of many curly multicelluar hairs.

* Rock cliffs in mountain valleys; 3500–4200 m. S Gansu.

14. Woodsia cinnamomea H. Christ, Bull. Acad. Int. Gιogr. Bot. 16: 122. 1906.

赤色岩蕨  chi se yan jue

Plants 8–15 cm. Rhizomes short, erect, scaly; scales brown, lanceolate, ca. 3 mm, membranous, entire, acuminate. Fronds densely clustered; stipe brownish stramineous, 2.5–5.5 cm, ca. 1 mm in diam., continuous, base densely scaly, upper part and rachis densely covered with long articulate hairs and brown linear scales, later mostly fallen. Lamina 1-pinnate or bipinnatifid, lanceolate, 6–8 Χ 1.5–2 cm, herbaceous, both surfaces densely covered with brown, articulate hairs, base slightly attenuate, apex acuminate. Pinnae in 1-pinnate lamina 9–11 pairs, sessile, well-separated, spreading; lower pairs slightly shortened; middle pinnae largest, oblong-ovate, 6–8 Χ 3–5 mm, close to rachises, base cuneate, margin undulate, apex obtuse; pinnae in bipinnatifid lamina ca. 7 pairs, opposite, obliquely spreading, 1 or 2 lower pairs somewhat shortened; middle ones longest, ovate, ca. 6 Χ ca. 4 mm, base broadly cuneate, pinnatifid, apex obtuse; segments hemielliptic, undulate. Veins obscure, pinnate, nearly reaching laminar margin. Sori round, consisting of a few sporangia, located on veinlets, medial. Indusia consisting of many curly, multicellular hairs.

* On rocks; ca. 2800 m. SW Sichuan (Muli).

15. Woodsia lanosa Hooker, Syn. Fil. 47. 1867 [??1866].

毛盖岩蕨  mao gai yan jue

Woodsia pellaeopsis Handel-Mazzetti.

Plants small, 2–10 cm. Rhizomes short, erect, scaly; scales brown, polished, linear, 2–3 mm, margin entire, apex filiform. Fronds clustered; stipe chestnut, slightly lustrous, 1–4 cm, less than 1 mm in diam., brittle, densely covered with brown, linear scales and articulate hairs. Lamina downward bipinnatifid, apically pinnatifid, brownish green when dried, lanceolate, 1–7 cm Χ 4–14 mm, herbaceous, densely covered with brown, curly, articulate hairs and linear scales abaxially, apex acuminate. Pinnae 4–10 pairs, sessile, well-separated, spreading; basal pair slightly shortened; middle ones equally large, narrowly elliptic, base truncate, pinnatifid, apex obtuse. Ultimate segments 2–4 pairs, hemielliptic, undulate. Veins obscure, pinnate, oblique, not reaching laminar margin. Sori round, terminal at vein tips, usually 2 or 3 per segment. Indusia hairlike, consisting of many curly, multicellular hairs.

On rocks in forests; 3100–4200 m. Sichuan, Xizang, Yunnan [India].

16. Woodsia andersonii (Beddome) H. Christ, Bull. Soc. Bot. France 52(Mιm. 1): 45. 1905.

蜘蛛岩蕨  zhi zhu yan jue

Gymnogramme andersonii Beddome, Ferns Brit. Ind. 2: 190, t. 190. 1866 [??not in Tropicos or IPNI].

Plants 10–20 cm. Rhizomes short, erect or ascending, scaly; scales dark brown, linear-lanceolate, ca. 3 mm, membranous, entire. Fronds clustered; stipe stramineous or brownish stramineous, lustrous, 5–10 cm, ca. 1 mm in diam., continuous, with fibrous small scales and articulate hairs, later partly fallen; rachis sparsely hairy when old, stipes and rachises persistent after pinnae fallen. Lamina basally 2-pinnatifid, apically pinnatifid, lanceolate, 5–10 Χ 1–2 cm herbaceous, both surfaces densely covered with rusty, articulate long hairs, especially when young, base not or slightly shortened, apex acuminate. Pinnae 6–9 pairs, sessile, lower pairs well-separated, not or slightly shortened; middle pinnae ovate or subrhombic, 7–12 mm, base broadly cuneate, lobed, apex obtuse. Ultimate segments 2–4 pairs, hemielliptic, entire or undulate. Veins obscure, pinnate, veinlets not reaching laminar margin. Sori round, located on acroscopic veinlets, 1–3 per segment, medial or supramedial. Indusia consisting of 8–10 long, curly hairs. 2n = 328.

Rock crevices and cliffs in forests; 2500–4500 m. Gansu, Qinghai, Shaanxi, Sichuan, Xizang, Yunnan [NE India (Sikkim)].

Woodsia andersonii is distributed in SW China and Himalayas. It is densely covered with rusty, long hairs when young, while most of the hairs fall away later.

17. Woodsia rosthorniana Diels in Engler, Bot. Jahrb. Syst. 29: 187. 1900.

密毛岩蕨  mi mao yan jue

Eriosoriopsis rosthorniana (Diels) Ching & S. H. Wu; Woodsia delavayi H. Christ; W. jeholensis Nakai & Kitagawa; W. lanosa Hooker var. attenuata C. Christensen.

Plants 7–25 cm. Rhizomes short, erect, scaly; scales brown, lustrous, linear-lanceolate, 4–5 mm, membranous, margin entire, apex subulate. Fronds clustered; stipe yellowish brown or stramineous, 2–6 cm, less than 1 mm in diam., densely covered with yellow-brown long multicellular hairs and linear scales, stipe and rachis usually curved and S-like; rachis as densely hairy and scaly as stipe. Lamina basally 1-pinnate-pinnatifid, apically pinnatifid, lanceolate, 7–20 Χ 1.5–3 cm, herbaceous, both surfaces densely covered with brownish long articulate hairs, base gradually shortened, apex acute. Pinnae 10–15 pairs, spreading, sessile, lower pairs reduced into auricles, well-separated; middle pinnae largest, oblong-elliptic or ovate-lanceolate, 8–15 Χ 5–8 mm, base broadly cuneate, pinnatifid, apex obtuse or rounded. Ultimate segments 4 or 5 pairs, hemielliptic, basal pair largest, entire. Veins obscure, pinnate, veinlets nearly reaching laminar margin. Sori round, located at vein endings or on veinlets, submarginal, dispersed throughout abaxial surface of pinnae at maturity. Indusia shallowly saucer-shaped, dissected into 3–5 plates at maturity, margin with curly, unicellular hairs. 2n = 164.

Rock crevices in forests and in thickets; 1000–3000 m. Hebei, Liaoning, Shaanxi, Sichuan, Xizang, Yunnan [Bhutan].

18. Woodsia cycloba Handel-Mazzetti, Symb. Sin. 6: 19. 1929 [??not in Tropicos or IPNI].

栗柄岩蕨  li bing yan jue

Plants 4–10 cm. Rhizomes ascending or prostrate, scaly; scales dark brown, lanceolate, ca. 2 mm, membranous, entire or sparsely dentate. Fronds nearly clustered; stipe castaneous, 2–5 cm, fragile, lower part covered with scales and articulate hairs, upper part and rachis densely covered with articulate knee-jointlike long hairs, hairs late fallen. Lamina imparipinnate, elliptic-lanceolate, 3–6 Χ 1–2.5 cm, thinly herbaceous, abaxially with articulate hairs along veins and margin, adaxially glabrous. Pinnae 5 or 6 pairs, opposite, spreading, sessile, ovate or elliptic, 8–12 Χ 5–9 mm, margin entire or slightly undulate, apex rounded. Veins visible, pinnate, veinlets not reaching laminar margin. Sori large, round, located at bifurcations of lateral veins or on acroscopic veinlets. Indusia absent or composed of ciliate platelike small lobes, hidden. 2n = 328.

Crevices in granite rocks and schist cliffs; 2900–4600 m. Shaanxi, Sichuan, Xizang, Yunnan [Nepal].

19. Woodsia okamotoi Tagawa, Acta Phytotax. Geobot. 7: 185. 1938.

岡本氏岩蕨  gang ben shi yan jue

Rhizomes ascending, up to 5 cm, densely covered by persistent stipe remains. Stipes clustered, rigid but fragile, together with rachis ebeneous, relatively polished, sparsely scaly and pubescent, 3–5 mm, up to 1 mm in diam.; hairs pale, articulate, flexuous; scales rust-colored, lanceolate or linear-lanceolate, membranous, gradually attenuate into articulate hairs, hairs not glandular-capitate at apex, scales in lower parts of stipe larger, up to 4 mm. Lamina 1-imparipinnate, lanceolate, widest at middle, 4–7 Χ 1–2 cm, herbaceous, slightly attenuate to base, apex obtuse. Pinnae 6–9 pairs, spreading, ± remote from each other, ovate or ovate-oblong, both surfaces pubescent, base truncate or finely cordate, sessile, crenate-lobed to pinnatifid, apex rounded; margin beneath to costa scaly; middle pinnae largest, 5–10 Χ 3–6 mm at base; lobes ca. 4 pairs, entire, rounded; terminal pinna smaller, 3–5 mm long and wide. Sori medial; indusia consisting of pale, long, crested hairs. Spores nearly ellipsoid, surface folded, tuberculate.

* Mossy rocks in coniferous forests. Taiwan.



[1] Herbarium, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China.

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

[3] University of Barnaul, Altai State University, Dimitrov’s Street 66, 656099 Barnaul, Russia.