ATHYRIACEAE (part 1) [FIRST DRAFT]
Wang Zhong-ren (王中仁)
ACYSTOPTERIS, CYSTOPTERIS, CYSTOATHYRIUM, GYMNOCARPIUM, ANISOCAMPIUM,
KUNIWATRSUKIA, PSEUDOCYSTOPTERIS, NEOATHYRIUM, ATHYRIUM, DRYOATHYRIUM,
LUNATHYRIUM
Subfam. 1. CYSTOPTERIOIDEAE
冷蕨亚科 leng jue ya ke
Medium- to small-sized plants, summer-green; rhizomes slender, creeping or ascending; laminae pinnate-pinnatifid to 3-(4-)pinnate-pinnatifid; veins free; sori small, round, dorsal on the veins; indusia ovato-lanceolate, ovate or round, attached at proximal side of sorophore; X=42.
Three genera included：ACYSTOPTERIS Nakai, CYSTOPTERIS Bernh., CYSTOATHYRIUM Ching; all of them occur in China.
1. ACYSTOPTERIS Nakai, Bot. Mag. Tokyo 47(555):180. 1933.
亮毛蕨属 liang mao jue shu
Cystopteris Luerss. non Bernh. 1806; Cystopteris subgenus Acystopteris (Nakai) Blasdell.
Medium-sized terrestrial plants. Rhizomes creeping, forked, dictyostele, sparsely scaly; scales lanceolate or ovato-lanceolate, lustrous, long-acuminate at apex, with sparse glandular-hair-like teeth at margin. Fronds approximate; stipes nearly as long as lamina, chestnut-brown or stramineous, with two vascular bundles (upwards united as "U" in transverse section of upper part of rachis), more or less bearing scales, hyaline multicellular hairs, and some scale-hairs (multicellular hairs, consist of two column cells at base, upward becoming one column), grooved adaxially; laminae broadly ovate to ovato-lanceolate, subacute to acuminate at apex, not narrowed toward base, 2- to 3-pinnate-pinnatifid; pinnae mostly subopposite, alternate only in the upper, broadly lanceolate to lanceolate, pinnatisect to 2-pinnate, patent or ascendant, slightly upwards curved at top, acuminate at apex, symmetrical, truncate or rotund-cuneate at base, subsessile or very shortly petiolulate, the basal pair of pinnae not shortened; pinnules ovato-lanceolate or lanceolate, sessile, obtuse or acuminate at apex, symmetrical, near truncate or rotund-cuneate at base, the basiscopic ones a little longer than the acroscopic ones, alternate, only the basal pair subopposite, a little shorter than the upper one; ultimate pinnules oblong, obtuse at apex, sessile or connate with costae or costules at base, lobed or parted at margin. Veins visible but not so clear, free, lateral veins pinnate, veinlets simple or forked, reaching tooth at margin. Laminae thin-herbaceous, surfaces and all costae and veins more or less bearing hyaline long multicellular hairs, costae more or less with scale-hairs too. Sori small, round, dorsal on veinlets, sorophore protuberant, two lines along two sides of costules; sporangium with two lines of alternate stalk cells, annulus cells of sporangium 12--14; indusia small, laterally symmetrical, membranous, pale-green, broadly ovate, glandular-hairy, sparsely ciliated at margin, attached at the proximal side of sorophore, hidden by sporangia at maturity; spores bean-shaped or long spheroidal, yellow, perispore surface densely with inflated, bulbous processes; X=42.
Three species occur in the world and in China. Of them, one is endemic in China. Occur in tropics and subtropics of SE Asia and New Zealand. China is as the distribution center, spreading East to Japan, West to India and Sikkim, South to Indo-China and Indonesia.
The genus ACYSTOPTERIS is very close to the genus CYSTOPTERIS. So it is often been treated as a subgenus of CYSTOPTERIS or simply put it in CYSTOPTERIS. However, the two genera occur in different geographic area and elevations. ACYSTOPTERIS occurs in tropical and subtropical region. But CYSTOPTERIS occurs in the temperate, cold-temperate zone and subtropical mountains. Leaves of ACYSTOPTERIS are larger, pinnae and pinnules are symmetrical at base, bearing hyaline multicellular long hairs and scale-like hairs all over. However, leaves of CYSTOPTERIS are rather smaller, pinnae and pinnules are usually asymmetrical at base, glabrous; the ornamentations of their spore surface are different. Therefore, ACYSTOPTERIS is kept as a separated genus here.
The Latin name of ACYSTOPTERIS is not keeping with the fact. Nakai, the nominator of the genus, misread its sori exindusiate when he published the new genus. In fact, all members of ACYSTOPTERIS are with small scale-like membranous indusia.
1a. Laminae broadly ovate to ovato-oblong; orange unicellular glands absent on the lower surface of lamina and at indusium margin; stipes and raches and costae chestnut-black or purple-brown; plants usually under 90 cm high; Provinces and regions in the South of the Yangtze River
1. A. japonica
1b. Laminae ovato-oblong to ovato-lanceolate; orange unicellular glands present on the lower surface of lamina and at indusium margin; stipes and raches stramineous or stipes and raches red-brownish, but costae stramineous; plant up to or more than 1 m high.
2a. Stipes and raches stramineous; multicellular hairs on the fronds dense; Taiwan and SW China
2. A. tenuisecta
2b. Stipes and raches brownish or dull brown; multicellular hairs on the fronds sparse. Taiwan
3. A. taiwainiana
1. Acystopteris japonica (Luerss.) Nakai in Bot. Mag. Tokyo 47(555):180. 1933.
亮毛蕨 liang mao jue
Cystopteris japonica Luerss. in Engler, Bot. Jahrb. 4:363. 1883.
Summer-green. Rhizomes creeping, 2--4 mm thick, sparsely clothed with yellowish-brown, broadly lanceolate, thin-membranous scales. Fronds approximate; fertile fronds (12--)40--60(--90) cm; stipes (6--)15--25(--40) cm, (1--)2--3(--4) mm thick, chestnut-black or purple-brown, grooved adaxially, sparsely bearing scales at bases, upwards glabrate, polished; laminae broadly ovate to deltoid-ovate, (6--)20--35(--50) cm, (3.5--)15--18(--40) cm broad, short-acuminate or acuminate at apex, subcordate at base, 2-pinnate to 3-pinnate; pinnae 10--15 pairs, the basal pair not shortened, oblong or broadly lanceolate, (1.8--)8--15(--25) cm long, (1--)3--6(--11) cm broad, acuminate at apex, subtruncate at base, patent or ascendant, subsessile or very shortly petiolulate, opposite or subopposite, 3--4 cm apart from the upper pair, pinnate-pinnatifid to 2-pinnate; pinnules 10--24 pairs, the basiscopic ones a little longer than acroscopic ones, the basal pair a little shortened, subopposite or anadromic, the second pair and the upwards oblong, (0.5--)2.5--4(--6) cm long, (3--)6--10(--20) mm broad, obtuse at apex, sessile, alternate, subsymmetrical, truncate or broadly cuneate at base, close to costae, pinnatifid to pinnate; segments 5--14 pairs, patent or slightly ascendant, oblong, obtuse at apex, crenate, cut at margin; the second pair of pinnae nearly same with the basal pair, but narrower, upper pinnae gradually shortened, upper pinnae lanceolate, pinnate or pinnatifid. Veins pinnated at segments, veinlets simple, extend to tooth. Laminae herbaceous, green when dried, along veins and costae, raches and stipes on both sides sparsely bearing hyaline multicellular long hairs, on the costae more or less mixed with scale-like hairs, raches with the same color as stipes, costae stramineous or only with the same color at base. Sori small, round, dorsal on the basal acroscopic veinlets of segments, uniserial along each side of midrib; indusia small, membranous, pale-green, broadly ovate, glandular and sparsely ciliated on margin, attached at the proximal side of sorophore, hidden by sori at maturity, look like inferior, persistent. Spores bean-shaped or long-spheroidal, yellow, perispore surface densely with inflated, bulbous processes. 2n=84@,168@.
In valley under forests; 400--2800 m. Zhejiang, Jiangxi, Fujian, Taiwan, Hubei, Hunan, Guangxi, Sichuan, Chongqing, Guizhou, Yunnan [Japan]. Type from Japan.
2. Acystopteris tenuisecta (Bl.) Tagawa in Acta Phytotax. Geobot. 7(2):73. 1938.
禾秆亮毛蕨 he gan liang mao jue
Aspidium tenuisectum Bl., Enum. Pl. Jav. 170. 1828 (Type K!); Alsophila tenuisecta Bl. ex T. Moore; Athyrium tenuisectum T. Moore; Asplenium tenuisectum Hook.; Cystopteris tenuisecta Mett.; Cornopteris tenuisecta Tard.-Blot; Lastrea setosa Bedd.; Cystopteris setosa Bedd.; Davallia setosa Bak.; Cystopteris formosana Hay.
Rhizomes creeping, 2--5 mm thick, sparsely clothed with brownish lanceolate thin-membranous scales. Fronds approximate; fertile fronds (22--) 65--80(--150) cm; stipes (7--)30--40(--90) cm long, (1.5--)3--4(--5) mm thick at base, pale stramineous, grooved adaxially, clothed with brownish lanceolate scales and sparse multicellular hairs at base; laminae ovate to ovato-lanceolate, (14--)35--50(--70) cm long, (7--)20--25(--60) cm broad, acuminate at apex, subcordate at base, 3-pinnate to 3-pinnate, ultimate pinnae pinnatisect; pinnae 15--20 pairs, nearly opposite, broadly lanceolate to lanceolate, acuminate at apex, slightly upwards curved, subsessile or shortly petiolulate, symmetrical, subtruncate at base, the basal 1--2 pairs of pinnae broadly lanceolate, (3.5--)15--20(--33) cm long, (2--)4--8(--20) cm broad, long acuminate at apex, subtruncate or rotund-cuneate at base, 2-pinnate to 2-pinnate-pinnatifid; pinnules 10--25 pairs, acroscopic one a little shorter than basiscopic one, the basal pair a little shortened, from the basiscopic second upwards pinnules broadly lanceolate, (1--)2.5--5(--10) cm long, (0.4--)1--2(--3) cm broad, obtuse at apex or acuminate at apex, truncate at base, sessile, alternate, pinnate to pinnate-pinnatifid; ultimate pinnules oblong, 4--10 mm long, 2--3 mm broad, obtuse at apex, sessile or more or less connate with costules at base, cut or lobed at margin; upper pinnae gradually shortened, similar to the basals or a little narrowed; terminal pinna lanceolate. Veins pinnate in segments, veinlets simple or forked. Laminae herbaceous when dried, pale-green, laminae on the abaxial surface and indusium margin bearing orange or yellow small glands, surfaces of veins, costules, costae, raches and stipes densely bearing hyaline multicellular long hairs; costae, raches same color with green stipes, bearing small subulate scales on the adaxial surface. Sori small, round, dorsal on the basal acroscopic veinlets of segments, uniserial along each side of costule; indusia small, membranous, pale-green, broadly ovate, attached at the proximal side of sorophore, hidden by sori at maturity, look like inferior, persistent. Spores bean-shaped or long-spheroidal, yellow, perispore surface densely with inflated, bulbous processes. 2n=84@,168@.
In valley under rain forests or damp area streamside in forests; 700--2600 m. Taiwan, Guangxi, Sichuan, Yunnan, Xizang [S Japan, Vietnam, Myanmar, Thailand, NE India, Nepal, Sikkim, Malaysia, Singapore, Indonesia, Philippines, New Zealand; the tropics of Asia]. Type from Indonesia (Java).
3. Acystopteris taiwaniana (Tagawa) Lφve et Lφve in Taxon 26(2--3):326. 1977 et in Lφve, Lφve et Pic. Ser., Cytotax. Atlas Pterid. 269. 1977.
台湾亮毛蕨 tai wan liang mao jue
Cystopteris japonica var. taiwaniana Tagawa in Acta Phytotax. Geobot. 4:55. 1935. (Type, KYO!, Isotype BM!); Acystopteris japonica var. taiwaniana (Tagawa) W. C. Shieh.
This species is in between Aystopteris japonica and A. tenuisecta morphologically. Stipes pale brown-red, costae stramineous, laminae bearing pale yellow glandular cells on the abaxial surface, spores normal. Diploid or tetraploid, 2n=84@,168@.
* Taiwan. Type from Ali Shan.
2. CYSTOPTERIS Bernh. in Schrad. Neu. Journ. Bot. 1(2):5,26. 1806.
冷蕨属 leng jue shu
FILIX Ludwig; CYSTEA Sm.; RHIZOMATOPTERIS A. P. Khokhr.
Small plants, summer-green. Rhizomes long- or short-creeping, dictyostele, blackish brown, glabrous or densely clothed with red-brown pubescence, sparsely scaly, scales brown or brownish, thin, ovate to broadly lanceolate; fronds distant, approximate or caespitose, thin. Stipes shorter or longer than lamina, dark brown at base, upwards stramineous or chestnut-coloured; laminae ovato-lanceolate, ovato-triangular or near pentagonal, 2-pinnate to 3-pinnate, rarely 4-pinnate or more divided; pinnae anadromic, shortly petiolulate, more or less unequal-sided or nearly symmetrical at base; small teeth present at segment margin; veins free, forked or pinnated, veinlets terminating in entire or emarginate teeth. Laminae thin-herbaceous or herbaceous, green when dried, stipes, raches and pinnae, pinnule sparsely bearing scale-hairs at base, multicellular hairs or unicellular glandular-hairs, laminae usually glabrous. Sori round, dorsal on veinlets, sorophore slightly protuberant, annulus of sporangium consists of 14-16 cells, indusia ovate, lanceolate, rotund or half cup-shaped, attached at the proximal side of sorophore under sori, membranous, persistent, covering sori when young, hidden by sori at maturity, look like inferior. Spores bilateral, bean-shaped, dark brown, perispore surface usually with spine-like processes, rarely with folds. X=42.
More than 20 species and many hybrids have been known in the world, mainly in the temperate zone, the frigid-temperate zone and tropical mountains under forests. 11 species in China, NE, N, NW, mountains of SW and Taiwan.
1a. Rhizomes short-creeping or ascending, fronds approximate or caespitose; laminae lanceolate to broadly lanceolate; stipes usually shorter than lamina, at most nearly as long as lamina; veins usually terminating in entire, rarely in emarginate teeth.
2a. Veins terminating in entire teeth.
3a. Perispore surface with densely spine-like processes; laminae usually narrower; on calcareous soil; Taiwan; NE, N, NW, mountains of SW China
1. C. fragilis
3b. Perispore surface rugose or verrucose without spine-like processes; laminae usually wider; on acid soil; Hebei; NW, SW China
2. C. dickieana
2b. Veins terminating in emarginate teeth.
4a. plant usually less than 10 cm high; stipes slender, a little longer than lamina or nearly as long as lamina, chestnut-coloured; Qinghai, Gansu, Sichuan, NW Yunnan and E Xizang
3. C. kansuana
4b. plant up to 29 cm high; stipes thick, ca. 1mm in diameter, shorter than lamina, dark purple-coloured at base; Guizhou
4. C. guizhouensis
1b. Rhizomes long-creeping, fronds distant; laminae broadly ovate, ovato-triangular or near pentagonal; stipes usually longer than lamina or as long as lamina; veins terminating in emarginate teeth.
5a. Laminae near pentagonal, the basal basiscopic pinnules of basal pair of pinnae prominently elongated.
6a. Ultimate segments not revolute at margin; spore surface with short spine-like or verrucose processes; N, NW, SW and Taiwan
5. C. montana
6b. Ultimate segments revolute at margin; spore surface with rugate folds; NW Yunnan
6. C. modesta
5b. Laminae broadly ovate or ovato-triangular, the basal basiscopic pinnules of basal pair of pinnae not elongated.
7a. Indusia glandular; plant thin; NE, N, Yunnan and Xizang
7. C. sudetica
7b. Indusia not glandular, plant moderately stout.
8a. Laminae bearing short glandular-hairs; E Xizang and NW Yunnan
8. C. tibetica
8b. Laminae lacking short glandular-hairs.
9a. Fronds up to 50 cm long or longer, laminae 4-pinnatipartite to 4-pinnatisect, perispore densely echinate; NW Yunnan
9. C. deqinensis
9b. Fronds less than 50 cm, laminae 2-pinnate to 3-pinnate, perispore sparsely echinate.
10a. Laminae 3-pinnate, ultimate pinnules or segments smaller and narrower, 3--5 mm broad; SW, Mt. Qinling, Hebei and Taiwan
10. C. moupinensis
10b. Laminae 2-pinnate to 2-pinnate, pinnules pinnatipartite, pinnules or segments bigger and wider, 5--7 mm broad; SW and Mt. Qinling
11. C. pellucida
1. Cystopteris fragilis (L.) Bernh. in Schard. Neu. Journ. Bot. 1(2):26--27. t. 2. f. 9. 1806.
冷蕨 leng jue
Polypodium fragile L. Sp. Pl. 2:1091. 1753; Cystopteris filix-fragilis (L.) Gilib.; Aspidium fragile Sw.; Athyrium fragile Spr.; Cyclopteris fragilis Gray; Cystea fragilis Sm.
Rhizomes short- 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; stipes usually shorter than lamina, nearly as 1/3--2/3 long as lamina, occasionally slender and a little longer than lamina when living in crevice, 5--14(--20) cm long, (0.2--)1--1.5 mm thick, brown at base, upwards stramineous or chestnut-coloured, bearing sparse, polished scales; laminae lanceolate to broadly lanceolate, 17--28 cm long, (0.8--)4--5(--8) cm broad, shortly acuminate at apex, usually 2-pinnate to 2-pinnate-pinnatifid, occasionally pinnate or 3-pinnate; pinnae 12--15 pairs, ascendant, the basal 1--2 pairs a little shortened, or nearly not shortened, ovate to ovato-lanceolate, (0.4--)2--4(--7) cm long, (0.2--)1--2.5 cm broad, obtuse or short-acuminate, toothed at apex, the acroscopic side parallel to raches, the basiscopic side more or less cuneate at base, subopposite, subsessile, apart from each other widely, usually 1-2 times the broad of pinnae, 1.5--4.5 cm; pinnules 5--7 pairs, ovate or oblong, rounded or obtuse and toothed at apex, the acroscopic side truncate, the basiscopic side cuneate at base, sessile or shortly petiolulate, entire or toothed at margin, or pinnatifid; the middle pinnae similar to the lower pinnae, but a little longer, separate from each other in 1.2--2.5 cm, subopposite or alternate, subsessile; upper pinnae pinnatipartite, toothed only on apices and acroscopic margin; veins pinnated, costules slightly tortuous, veinlets terminating in entire teeth. Laminae herbaceous when dried, green or yellow-green. Raches and costae, especially on the base more or less bearing sparse unicellular to long multicellular hairs, even a few scale hair. Sori small, round, medial on veinlets, 2--4 pairs per pinnule, 1--2 sori at acroscopic side in upper pinnules; indusia ovate to lanceolate, membranous, pale-green or brownish. Spores dark-brown, perispore surface with regular, denser spine-like processes. 2n=84@,168,252,336.
Under alpine shrubs, rock crevice on shady slope, rock foot or damp area streamside; (200--)1500--4500(--4800) m. Heilongjiang, Jilin, Liaoning, Nei Mongol, Hebei, Shanxi, Shaanxi, Ningxia, Gansu, Qinghai, Xinjiang, Shandong, Anhui, Taiwan, Henan, W Sichuan, Yunnan, Xizang [Korea, Japan, Mongolia, Russia, India, Nepal, Pakistan, Afghanistan, Iran, Turkey; Europe, the North, central mountains and Himalayas of Asia, North America, South America, Africa]. Type from Europe.
2. Cystopteris dickieana R. Sim, Gard. Farmer's Journ. 2:308. 1848.
皱孢冷蕨 zhou bao leng jue
Cystopteris fragilis var. dickieana (Sim) Lindberg in Medd. Soc. Fauna et Fl. Fenni32:21--24. 1905.; Cystopteris fragilis subsp. dickieana (Sim) Hylander; Cystopteris baenitzii Dorfl.; Cystopteris sikkimensis Ching ex Bir.
Very similar to Cystopteris fragilis, hard to be distinguished from the later morphologically except spores ornamentation. Perispore surface 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.
In China, common in Xizang and Xinjiang, Sympatric in the distribution area of C. fragilis.
In valley or in rock crevice on mountain slope, on rocks under forests, and grassland damp areas; 1400--5400(--5600)m. Hebei, Shaanxi, Gansu, Qinghai, Xinjiang, Taiwan, Sichuan, Yunnan, Xizang [India, Nepal, Pakistan, Afghanistan; Europe, the North and central mountains of Asia, North America, Africa]. Type from Europe (Scotland).
The taxonomic status of the species is uncertain. At present, we know the Chinese C. dickieana only based on the spore surface which is not echinate. However, their spore surface ornamentation is different in the specimens from different localities. Their morphology and habitat also are different from the type of C. dickieana. Are the Chinese plants another species or the same species with European specimens? That is still a question.
3. Cystopteris kansuana C. Chr. in Journ. Wash. Acad. Sci. 17:499. 1927 (Isotype: PE!).
西宁冷蕨 xi ning leng jue
Cystopteris X kansuana Blasdell.
Rhizomes slender, short-creeping, sparsely scaly. Fronds approximate; fertile fronds 10--19 cm; stipes 6--11 cm long, less than 1 mm thick, chestnut-coloured, bearing a few of brown lanceolate scales at base, upwards glabrate; laminae lanceolate, 4--10 cm long, 1.3--3 cm broad, long-acuminate at apex, obtuse at base, pinnate, pinnae pinnatipartite to pinnatisect; pinnae 5--8 pairs, the basals subopposite, upward alternate, ascendant, subsessile or shortly (1--2 mm) stalked, basal pair of pinnae a little shortened, ovate to long ovate, 9--20 mm long, 4--8 mm broad, acuminate at apex, basal acroscopic broadly cuneate, the basiscopic side narrowly cuneate, pinnatipartite to pinnatisect; segments ca. 3 pairs, ascendant, the basal acroscopic one bigger, long ovate, 4--8 mm long, 2--3 mm broad, acute or obtuse-rounded at apex, crenate or dentate-pinnatipartite at margin, the rest segments oblong or linear-oblong, far smaller, acuminate at apex, with 2--3 small teeth, entire at margin, segments joining each other by narrow wing at base; the second pair of pinnae 1--2.4 cm apart from the basal pair of pinnae, similar and a little bigger than the basal pair, upward the pinnae gradually smaller. The first basal acroscopic lateral veins of segments pinnated, the rest lateral veins simple or occasionally forked, terminating in emarginate teeth. Laminae pale-green or dark brown when dried, thin-herbaceous. Sori round, yellow-brown, medial on veinlets, indusia ovate, entire at margin, pale yellow-brown, persistent. Spore perispore surface echinate (not from type specimen).
* In shaded rock crevice; 3000--4500 m. Gansu, Qinghai, Sichuan, Yunnan, Xizang. Type from Qinghai (Xining).
The reason of its Latin name being named as "kansuana" was that Xining was subordinated to Gansu Province then.
4. Cystopteris guizhouensis X. Y. Wang et P. S. Wang in Acta Bot. Yunnan. 19(2):141--142, f. 1. 1997. (Type HGAS!)
贵州冷蕨 gui zhou leng jue
Rhizomes short-creeping, scaly at apex and stipe bases, scales brown, ovato-lanceolate. Fronds approximate or caespitose; fronds up to 29 cm; stipes longer than laminae, 7--9 cm, nearly as 1/2 long as laminae, dark purple at base, upwards stramineous; laminae lanceolate, 18--21 cm long, 4--5 cm broad at middle, long-acuminate at apex, slightly narrowed at base, pinnate; pinnae 12--15 pairs, the basals subopposite, sessile, patent, basal1--2 pairs a little shortened, oblong or ovato-oblong, 2.5--2.8 cm long, 1--1.2 cm broad, obtuse at apex, nearly symmetrical, cuneate at base; segments 8--10 pairs, often oblong, rounded or truncate and crenate at apex; veins free, veinlets terminating in emarginate teeth. Laminae thin-herbaceous when dried, dark green, glabrous on surfaces. Sori small, round, usually only one row between midrib and margin in a pinna; indusia broadly ovate, membranous, persistent. Perispore surface more dense spine-like ornamentation, spine-like processes usually long 6--7μｍ.
* Under forests, rock slits; ca. 2800 m. Guizhou. Type from Guizhou (Hezhang).
5. Cystopteris montana (Lam.) Bernh. ex Desv. in Journ. Bot. Schrad. 1(2):26. 1806.
高山冷蕨 gao shan leng jue
Polypodium montanum Lam. Fl. Franc. 1:23. 1778; Aspidium montanum Sw.; Athyrium montanum Rohl ex Spr.; Rhzomatopteris montana A. P. Rhokhr.
Rhizomes long-creeping, blackish brown, glabrous, sparsely clothed with brownish ovate membranous scales, more densely scaly at apex. Fronds distant; fertile fronds 20--49 cm; stipes (6--)14--31 cm, as 1--3 times long as lamina, blackish brown at lower part, sparsely bearing brownish ovate scales at base, upwards stramineous or pale chestnut-coloured, glabrous, with a few scales; laminae near pentagonal, (5--)8--15(--20) cm, broad and long nearly equal, acuminate at apex, 3- to 4-pinnate, rarely up to 4-pinnate-pinnatifid; pinnae 4--7(--10) pairs, the basals subopposite, upward gradually alternate, patent, shortly petioled, the basal pair largest, deltoid-ovate or triangular, (2.5--)6--11(--13) cm long, (2--)4--7 cm broad, petioles 3--10 cm, acuminate at apex, broader, subtruncate, asymmetrical at base, 2-pinnate, ultimate pinnules pinnatipartite; pinnules (3--)6--8(--10) pairs, anadromic, acroscopic pinnules triangular, (0.4--)1--2 cm long, (6--)8--9(--12) mm broad, acuminate at apex, near truncate, symmetrical at base, the basiscopic pinnules prominently longer than the acroscopic ones, nearly two times long, the basal basiscopic pinnules of the lowest pinnae largest, oblong-triangular, as 2--3 times long as the acroscopic ones, 3--5 cm long, 1.5--2.5 cm broad, exserted downward nearly in right angle, sessile or with 1--2 mm short petiolule, acuminate at apex, truncate at base; secondary pinnules ca. 6 pairs, ovate to oblong, alternate, patent, near acute at apex, sessile, often decurrent and connate with costules at base, secondary pinnules the acroscopic one shorter, deltoid-ovate, 4--9 mm long, 3--6 mm broad, basiscopic one longer, up to 8--16 mm long, 6--9 mm broad; tertiary pinnules 4--5 pairs, obtuse-rounded at apex, cuneate at base, subopposite, ascendant, joining by narrow wing at base, up to pinnatifid; ultimate segments ovate or broadly lanceolate, rounded-obtuse at apex, subopposite, ascendant, joining by narrow wing at base, in some case pinnatifid, segments slightly incised at apex, entire at margin; the second pair of pinnae ca. 2 cm apart from the basal pair of pinnae, up to 3.5 cm long, similar to the lowest pinnae but smaller, upper pinnae gradually smaller, acroscopic and basiscopic pinnules gradually become equal. Veins pinnate, costules slightly tortuous, veinlets simple or forked, terminating in emarginate teeth. Laminae thin-herbaceous when dried, greenish brown, raches, costae and costules more or less bearing unicellular to multicellular hyaline hairs or short glandular-hairs. Sori small, round, brown, dorsal on veinlets, 3-7 sori per ultimate segment, 1 sorus per lobe; indusia rotund, pale-green or yellow-brown, thin-membranous. Perispore surface with short spine-like or verrucose processes. 2n=168.
Alpine mountains, wet areas under forests; 1700--4500 m. Nei Mongol, Hebei, Shanxi, Shaanxi, Ningxia, Gansu, Qinghai, Xinjiang, Taiwan, Henan, Sichuan, Yunnan, Xizang [E Europe, Korea, Japan, Russia, N India, E Pakistan; North America. Type from Europe.
6. Cystopteris modesta Ching in Bull. Fan Mem. Inst. Biol. Bot. 10:5. 1940 (Type PE!).
卷叶冷蕨 juan ye leng jue
Cystopteris sudetica var. moupinensis Blasdell., pro parte.
Rhizomes long-creeping, blackish brown, glabrous, sparsely clothed with brownish broadly ovate membranous scales, more densely scaly at apex. Fronds distant; fertile fronds 10--30 cm; stipes 6--17 cm, brown or chestnut-coloured at base, sparsely bearing brownish ovate scales, upwards stramineous and with only a few scales; laminae ovate, 5--13 cm long, 4--7 cm broad, acuminate at apex, broadly cuneate at base, 3-pinnate, ultimate pinnules pinnatipartite; pinnae 4--5 pairs, alternate, upswept, petiole 3--5 mm, the basal pair the largest, narrowly deltoid-ovate, ca. 7 cm long, ca. 2.5 cm broad, acuminate at apex, broadly cuneate at base, 2-pinnate; pinnules 5--9 pairs, ca. 8 mm apart from each other, oblong-ovate, acuminate at apex, near truncate, symmetrical at base, shortly petiolulate, alternate, very upswept, the basal basiscopic pinnules of lowest pinnae the longest, up to 3.2 cm, acroscopic and basiscopic pinnules gradually become equal upward; secondary pinnules 4--5 pairs, oblong, 3--6 mm, gradually narrowed at apex, obtuse at apex, subsessile, not decurrent and separate from costules at base, pinnatifid; ultimate segments small, 2--3 pairs, close, broadly ovate, rounded-obtuse at apex, decurrent to narrow costal wing at base, revolute at margin, subentire; the second pair of pinnae is ca. 3 cm apart from the basal pair, similar to the basal pinnae but smaller; upper pinnae gradually smaller. Veins pinnate in segments, veinlets free. Laminae greenish brown when dried; raches, costae and midribs glabrate. Sori round, dorsal on veinlets; indusia brown, persistent. Perispore surface with rugate folds, clearly with foveolate or reticulate ornamentation on the surface.
* In rock crevice streamside; ca. 3600 m. NW Yunnan (Watershed between Nujiang River and Qiujiang River).
Very similar to Cystopteris montana, but ultimate segments revolute at margin, spore surface without short spine-like or verrucose processes, with rugate folds, clearly foveolate on the surface, in some case, finely reticulate, easy to be recognized.
7. Cystopteris sudetica A. Br. et Milde, 33 Jahresber. Schles. Gesellsch. Vaterl. Kult. 92. 1855.
欧洲冷蕨 ou zhou leng jue
Rhizomatopteris sudetica A. P. Khokhr.; Cystopteris leucosoria Schur.
Rhizomes long-creeping, 1--2 mm thick, rhizome and stipe bases clothed with brown short hairs and a few pale-brown membranous ovato-lanceolate scales, more densely scaly at apex. Fronds distant; fertile fronds (15--)20--30 cm; stipes 10--16(--20) cm, thin, stramineous, polished; laminae broadly ovate or ovato-triangular, 9--15(--20) cm long, 8--12(--15) cm broad, acuminate at apex, 3-pinnate; pinnae 8--12 pairs, ascendant, the basal pair not shortened, oblong or ovato-lanceolate, (3--)4.5--7(--8) cm long, 1.8--3 cm broad at middle, acuminate at apex, slightly narrowed, asymmetrical at base, with 2--3 mm short petiolules, subopposite, (1--)1.5--2.5(--3) cm apart from the second pair of pinnae, 2-pinnate; pinnules 8--12 pairs, anadromic, acroscopic one a little shorter than basiscopic one or nearly equal, the second basiscopic one the largest, ovate or ovato-triangular, 1--2 cm long, 5--8 mm broad, obtuse or acute at apex, toothed, unequal-sided at base, the acroscopic side parallel to costae, the basiscopic side cuneate, subsessile or with 1--2 mm short petiolule, alternate, pinnate; secondary pinnules or segments 4--5 pairs, broadly ovate to oblong, the basal acroscopic one largest, 5--6 mm long, 3--4 mm broad, rounded-obtuse or subtruncate and serrate at apex, broadly cuneate, free or connate with costules by narrow wing at base, cut or toothed at margin; the rest pinnules upward gradually smaller, oblong-obovate, crenate at apex, the basiscopic side entire, the acroscopic side with 1-2 lobes; the second pair of pinnae similar to the basal pair, slightly shortened, subopposite, 1--1.6(--2) cm apart from the basal pair; upper pinnae lanceolate, pinnatifid, segments serrate at apex, toward base entire at margin. Veins visible on surfaces, veinlets simple or 1--2 times forked, terminating in emarginate teeth. Laminae thin-herbaceous or herbaceous when dried, green. Raches and costae occasionally with sparse or dense short glandular-hairs and a few multicellular long hairs. Sori small, rotund, 1-2 sori in ultimate pinnae or segments, often dorsal on suprabasal veinlets, sori more than 2 in the basal acroscopic bigger pinnules of pinnae, biserial along two sides of costules; indusia near round or half-cup shaped, pale-brown or yellow-brown, with sparse tiny glandular hairs. Perispore surface with spine-like processes. 2n=84,168.
Under coniferous or mixed forests; 900--3300 m. Heilongjiang, Jilin, Liaoning, Nei Mongol, Hebei, Shanxi, Yunnan, Xizang [Japan, Korea, E Russia, Europe]. Type from Europe.
8. Cystopteris tibetica Z. R. Wang in Acta Phytotax. Sin. 32(1):85, t. 1, f. 7. 1994 (Holotype PE!).
藏冷蕨 zang leng jue
Rhizomes long-creeping, 1.5--2 mm thick, rhizome and stipe bases clothed with brown hairs and a few membranous scales. Fronds distant; fertile fronds 15--20 cm; stipes 6--14 cm long, 1--1.5 mm thick, stramineous, at base brownish, clothed with brown hairs and a few broadly ovate or ovato-lanceolate scales; laminae deltoid-ovate, 6--11 cm long, 3--5 mm broad, acuminate at apex, 3-pinnate; pinnae 8--10 pairs, alternate, the basal pair subopposite, upswept, the basal pair the largest, oblong or ovato-lanceolate, 3--5 cm long, 1.2--1.5 cm broad, acuminate at apex, slightly narrowed and asymmetrical at base, petioles 1.5--3 mm, 2-pinnate, 1.2--3 cm apart from the second pair of pinnae, the second pair of pinnae same as the basal pair, a little small, top pinnae lanceolate; pinnules 6--8 pairs, anadromic, usually acroscopic one a little shorter than basiscopic one, the second basiscopic pinnules the largest, deltoid-ovate, 6--8 mm long, 4--5 mm broad, obtuse at apex, unequal-sided at base, the acroscopic side parallel to costae, the basiscopic side broadly cuneate, toothed at margin, very shortly petiolulate, alternate, pinnatilobate to pinnatisect; ultimate pinnules or segments ca. 3 pairs, the basal acroscopic the largest, ovate, oblong or obovate, 2--3 cm long, 1.5--2 mm broad, obtuse at apex, cuneate and connate with wings of costules or partly separate at base, serrate at margin, upper ultimate segments rhomboid or short-linear, only the acroscopic side and apices teethed. Veins visible on surfaces, veinlets pinnate to several times pinnate, terminating in emarginate teeth. Fronds thin-herbaceous when dried, green, clothed with short glandular-hairs on surfaces. Sori small, round, 1-2 sori per ultimate segment, dorsal on suprabasal veinlets, often 3-4 in the basal acroscopic pinnules; indusia rotund or half-cup shaped, pale-brown, frequently bearing a few glands, covering sori when young, hidden by sporangia at maturity.
On alpine slope, damp areas under coniferous forests, streamside, on rocks or tree trunks; 2400--3600 m. Yunnan, Xizang [Sikkim]. Type from Xizang (Mainling).
Similar to Cystopteris moupinensis, but lamina bearing short glandular-hairs on surfaces, indusia with short glandular-hairs too.
9. Cystopteris deqinensis Z. R. Wang in Acta Phytotax. Sin. 32(1):84, t.1, f.6. 1994 (Holotype PE!).
德钦冷蕨 de qin leng jue
Rhizomes long-creeping. Fronds distant; fertile fronds 45--60 cm; stipes 20--25 cm long, 1.5--3 mm in thick, stramineous, dark-brownish at base, clothed with brown hairs and a few broadly ovate or lanceolate scales; laminae narrowly ovate or ovato-oblong, 25--35 cm long, 15--20 cm broad, acuminate at apex, 3-pinnate, ultimate pinnules pinnatipartite, nearly pinnatisect; pinnae up to 10--13 pairs or more, alternate, the basal pair the largest, subopposite, upswept, ovato-lanceolate, 12--17 cm long, 4--6 cm broad, acuminate at apex, slightly narrowed and asymmetrical at base, petiole 0.5--1.5 cm, 5--7 cm apart from the second pair of pinnae; pinnules 10--15 pairs, anadromic, usually basiscopic one a little longer than acroscopic one, especially in the basal pair of pinnae, its basiscopic second and third pinnules largest, ovate to broadly lanceolate, 2.5--4.3 cm long, 1.5--2.2 cm broad, acuminate at apex, unequal-sided at base, the acroscopic side parallel to costae, the basiscopic side broadly cuneate, petiolules 1.5--2 mm, alternate; the second pair of pinnae almost same with the first pair in size, upper pinnae gradually smaller; ultimate pinnules 5--7 pairs, the basal acroscopic one the largest, broadly ovate, 8--14 mm long, 5--8 mm broad, rounded-obtuse at apex, narrowly cuneate at base, pinnatifid, toothed at margin. Veins visible on surfaces, veinlets forked, sometimes pinnate to 2-pinnate, veinlets terminating in emarginate teeth. Laminae thin-herbaceous when dried, green. Sori round, 1-6 sori per ultimate segment, dorsal on veinlets; indusia rotund or half-cup shaped, pale-green to brownish, without glands, covering sori when young, hidden by sporangia at maturity. Spores bean-shaped, perispore surface with very densely spine-like processes.
* In valley on rocks or under oak forests on mountain slope, streamside; 2400--3000 m. NW Yunnan. Type from Yunnan (Deqen).
This species look like Cystopteris moupinensis, but plant larger, fronds up to 60 cm, laminae narrowly ovate or ovato-oblong, 3-pinnate, ultimate pinnules pinnatipartite, spore surface with densely spine-like processes.
10. Cystopteris moupinensis Franch. in Nouv. Arch. Mus. Hist. Hat. Paris ser. 2, 10:111. 1887 (Type P!).
宝兴冷蕨 bao xing leng jue
Cystopteris sudetica var. moupinensis C. Chr.; Cystopteris sphaerocarpa Hay.; Cystopteris mairei Brause; Cystopteris tangutica Grubov; Cystopteris sudetica auct., non A. Br. et Milde. 1855.
Rhizomes long-creeping, 1--2 mm thick, rhizomes and stipe bases clothed with brown hairs and a few pale-brown broadly ovate membranous scales. Fronds distant; fertile fronds (20--)30--40(--50) cm; stipes 10--22(--25) cm long, 1--2 mm thick, stramineous or chestnut-brown, polished; laminae ovate or deltoid-ovate, 9--17(--25) cm long, 5--8(--15) cm broad, acuminate at apex, pinnate-pinnatifid to 3-pinnate; pinnae 8--12(--15) pairs, upswept, the basal pair oblong or ovato-lanceolate, 5--10 cm long, 2--3.5 cm broad, acuminate at apex, toothed, slightly narrowed and asymmetrical at base, petiolule 3--4 mm, subopposite, 3--3.5(--5) cm apart from the second pair of pinnae, 2-pinnate; pinnules 8--12 pairs, anadromic, usually acroscopic one a little shorter than basiscopic one, the second basiscopic the largest, deltoid-ovate or long triangular, 1--2.2(--3) cm long, 8--13 mm broad, obtuse and toothed, unequal-sided at base, the acroscopic side parallel to costae, the basiscopic side broadly cuneate, petiolule 1--2 mm, alternate, pinnate or pinnatipartite; ultimate pinnules or segments 3--4 pairs, the basal acroscopic one the largest, broadly ovate, 4--9 mm long, 3--5 mm broad, rounded-obtuse and toothed at apex, cuneate and connate with wings of costules at base, cut or toothed at margin, upper ultimate pinnules rhomboid or short-linear, apex and acroscopic toothed at apex; the second pair of pinnae same as the basal pair but a little shortened and narrowed, petiole 2--3 mm, 1.5--3 cm apart from the upper pair; the top pinnae lanceolate, pinnatifid, segments only with 1 or 2 teeth at apex. Veins visible on surfaces, veinlets one to several times forked, terminating in emarginate teeth. Laminae thin-herbaceous when dried, green. Sori small, round, dorsal on suprabasal veinlets, 1--2 sori per ultimate segment, in the acroscopic bigger pinnules of pinnae often 3--4 or more, biserial along two sides of costules on veinlets; indusia rotund or half-cup shaped, pale-green or yellow-brown, membranous, without tiny glandular hairs, covering sori when young, hidden by sporangia at maturity. perispore surface bearing spine-like processes.
Under mixed forests, damp areas or on wet rocks; 1000--4100 m. Hebei, Shaanxi, Gansu, Qinghai, Taiwan, Henan, Sichuan, NW Guizhou, Yunnan, SE Xizang [Japan, Sikkim and N India]. Type from Sichuan (Baoxing).
Very similar to Cystopteris sudetica, the main difference is that indusia without tiny glandular hairs, plant slightly moderately stout; phytogeographically C. sudetica occur in NE and N China, occasionally in high mountains of Yunnan and Xizang. But C. moupinensis mainly occur in mountains of W China, north up to Mt. Qinling, west up to E Xizang.
11. Cystopteris pellucida (Franch.) Ching ex C. Chr., Ind. Fil. Suppl. 3:67. 1934.
膜叶冷蕨 mo ye leng jue
Aspidium pellucidum Franch. in Nouv. Arch. Mus. Hist. Hat. Paris Ser. 2, 10:119. 1887. (Type P!); Nephrodium pellucidum Diels; Dryopteris pellucida C. Chr.; Cystopteris alata Ching.
Rhizomes long-creeping, ca. 2 mm thick, rhizome and stipe bases densely clothed with red-brown short hairs, and a few pale-brown broadly ovate membranous scales. Fronds distant; fertile fronds (20--)50--60 cm; stipes (10--)20--32 cm long, 1--2 mm thick, pale stramineous or red-brownish, polished, bearing a few pale-brown, broadly ovate, membranous scales mixed with short hairs at base; laminae ovate to narrowly ovato-oblong, (10--)20--33 cm long, (5--)10--15(--25) cm broad, long-acuminate at apex, pinnate, pinnae pinnatifid to 2-pinnate-pinnatifid; pinnae (10--)12--15(--17) pairs, upswept, the basal pair deltoid-lanceolate, (5--)8--14 cm long, 2.5--4.5 cm broad, caudate-acuminate at apex, asymmetrical at base, petiole 3--5(--8) mm, subopposite, (3--)5(--7) cm apart from the second pair of pinnae, pinnate, pinnules pinnatipartite; pinnules 8--12 pairs, anadromic, alternate, usually acroscopic one a little shorter than basiscopic one, the second basiscopic basal pinnae the largest, long-ovate or long-triangular, 1.5--2.5(--4) cm long, 1--1.5 cm broad, acute or near obtuse and toothed at apex, very asymmetrical at base, the acroscopic side narrowly cuneate, and parallel to costae, abaxial side broadly cuneate, upswept, very shortly petiolulate or subsessile, pinnatipartite; segments 3--5 pairs, the basal acroscopic one largest, oblong or ovate, rounded-obtuse or subtruncate at apex, toothed at apex and margin; pinnae from to the sinus or incision between teeth. The second pair gradually shortened upward, usually the basal acroscopic pinnules larger, triangular, segments toothed only at apex, entire at margin. Veins visible on surfaces, clearer abaxially, veinlets simple or forked 1--2 times, terminating in emarginate teeth. Laminae thin-herbaceous or near membranous when dried, green. Sori round, brown, dorsal on suprabasal veinlets, 1-2 sori per ultimate segment, 3-4 or more in the basal acroscopic larger pinnules of pinnae, biserial along two sides of costules on veinlets; indusia rotund or half cup-shaped, pale-brown, membranous, without tiny glandular hairs, covering sori when young, hidden by sporangia at maturity. Perispore surface bearing spine-like processes. 2n=ca.84.
* On mountain slopes under forests or damp area streamside; 1500--3700 m. Shaanxi, S Gansu, Henan, W Sichuan, Yunnan, Xizang. Type from Sichuan (Baoxing).
Very similar to and sympatric with Cystopteris moupinensis, different from the later in plant a little bigger and more stout, laminae division a little lower, but longer and narrower, ultimate segments wider, of thinner texture. They are hard to be distinguished in some degree. The relationship between them is still waiting for further studying.
3. CYSTOATHYRIUM Ching in Acta Phytotax. Sin. 11:22. 1966.
光叶蕨属 guang ye jue shu
ATHYRIUM Kato et Kramer, pro parte.
Medium-sized plant, evergreen, under forest, at moist place. Rhizomes ascending, bearing remainder stipe bases and dense thick roots, clothed with dark-brown ovato-lanceolate scales at apex. Fronds approximate; stipes short, brown and a little swollen at base, clothed with a few scales, upwards pale stramineous, glabrate, grooved adaxially; laminae narrowly lanceolate, near papery, gradually narrowed towards two ends, pinnate-pinnatifid, with scattered orange glands on the abaxial surface; pinnae numerous, up to 30 pairs, sessile, falcate-lanceolate, acuminate at apex, somewhat asymmetrical at base, lower pinnae gradually shortened towards the base, pinnatipartite cut to the wings of costae; segments oblong, obtuse at apex, upswept, entire or sparsely crenate at margin; costae grooved adaxially; veins pinnate in segments, visible on surfaces, a little protuberant adaxially, lateral veins 3--5 pairs, upswept, anadromic, simple, basal veins of basal segments frequently forked, reach to the margin of the leaves. 1--3 unknown small processes present on the abaxial surface of simple lateral veins in the basiscopic side of costa at the most middle pinna apices. Sori round, 1 sorus per segment, dorsal on the basal acroscopic veinlets, uniserial along each side of costule; indusia broadly ovate, thin-membranous, inferior (i.e. hidden by sporangia) at base, partly covering sori when young, hidden by sporangia at maturity, persistent. Annulus consisted of 1213 cells. Spores bean-shaped, dark-brown, darkened, perispore surface with dense tapered spine-like processes.
One species. W Sichuan (Erlang Shan, Tianquan).
This genus is taxonomically in between ATHYRIUM Roth. and CYSTOPTERIS Bernh. It look like the former, but its rhizome short-ascending, glabrate, stipe not beak-shaped at base, spore surface with spine-like processes, indusia inferior which are like the latter; However, it is different from the latter in lamina narrowly lanceolate, near papery, evergreen, stipe short, gradually narrowed towards the base, pinnate, pinnae pinnatipartite, pinnae much more, sessile, falcate-lanceolate. The special character different from the both genera is that there are some unknown small processes present on the basiscopic side of the middle pinna at apices.
1． Cystoathyrium chinense Ching in Acta Phytotax. Sin. 11:23, t. 4. 1966 (Type PE!).
光叶蕨 guang ye jue
Rhizomes short, ascending, bearing remainder stipe bases, clothed with brownish ovato-lanceolate scales at apex. Fronds approximate; fertile fronds 40--45 cm; stipes up to 7--8 cm long, ca. 2 mm thick, brown and a little swollen at base, slightly bearing 1 or 2 lanceolate scales, upwards stramineous, glabrate, grooved adaxially; laminae narrowly lanceolate, up to 35 cm long, 6--8 cm broad at middle, gradually narrowed towards two ends, pinnatifid-acuminate at apex, lower pinnate, pinnae pinnatipartite; pinnae ca. 30 pairs, subopposite, patent, sessile, ca. 1 cm apart from each other (the lowers wider apart), the basal pair only ca. 1 cm, triangular, the longest pinnae at middle 3--4 cm long, ca. 1 cm broad at base, narrowly falcate-lanceolate, acuminate and curved at apex, asymmetrical at base (the acroscopic side broader, truncate, the basiscopic side narrower-cuneate or obtuse-rounded), pinnatipartite cut to narrow wings of costae; segments up to ca.10 pairs, upswept, oblong, obtuse at apex, separated by narrow incisions, acroscopic basal segments a little longer than basiscopic one, the basal two segments the largest, 5--8 mm long, ca. 3 mm broad, gradually shortened upwards, the basal basiscopic one near broadly ovate, a little shortened, entire at margin, or basal 1--2 pairs slightly crenate. Veins in segments pinnate, lateral veins anadromic, 3--5 pairs, simple, oblique, terminating in to the margin of leaves. Laminae near papery when dried, pale-green, glabrous; raches grooved adaxially, glabrous. Sori round, 1 sorus per segment, dorsal on the basal acroscopic veinlets, close to costae, uniserial along each side of midrib; indusia round, thin-membranous, pale-green, fugacious, look like exindusiate. Spores bean-shaped, dark-brown, darkened, perispore surface densely spinate.
* Damp areas under forests; 2400--2500 m. W Sichuan. Type from Sichuan (Erlang Shan, Tianquan).
Subfam. 2. ATHYRIOIDEAE
蹄盖蕨亚科 ti gai jue ya ke
Plants small to large; rhizomes erect, ascending or creeping; laminae pinnate to 3-pinnate-pinnatifid; sori usually horseshoe-shaped, J-shaped, short-linear, lunate, reniform or oblong etc., in some cases, various forms of sori exist in a same plant, even in a same lamina or segment, cross over, lateral or dorsal on veinlet; indusia same with sori in shape; veins free, occasionally reticular; basal chromosome number X=40.
10 genera included：GYMNOCARPIUM Newman, KUNIWATRSUKIA Pic.-Serm., ANIDOCAMPIUM Presl, PSEUDOCYSTOPTERIS Ching, DRYOATHYRIUM Ching, LUNATHYRIUM Koidz., ATHYRIOPSIS Ching, ATHYRIUM Roth, NEOATHYRIUM Ching et Z. R. Wang, and RHACHIDOSORUS Ching.
4. GYMNOCARPIUM Newman in Phytologist 4:371. 1851.
羽节蕨属 yu jie jue shu
CURRANIA Cop.; CARPOGYMIA Lφve et Lφve.
Medium- to small-sized plant, terrestrial, Summer-green. Rhizomes long-creeping, blackish brown, glabrate, dictyostele, clothed with brown, thin, broadly lanceolate or ovato-lanceolate scales at apices and stipe bases. Fronds distant; stipes thin, far longer than lamina, dark brown at base, stramineous upwards, U-shaped grooved adaxially; laminae deltoid-ovate to pentagonal-oval, acuminate at apex, bases articulate to the stipe apex, simple-pinnatipartite to 3-pinnate-pinnatifid; pinnae petioled or sessile, articulate to raches, the basal pair not shortened. Veins free, pinnate in ultimate segments, lateral veins simple or occasionally forked, terminating to margin. Laminae herbaceous or thin-herbaceous, stipes top, raches, costae and lamina more or less bearing hyaline or pale yellow glands on surfaces, or absent. Sori oblong or round, exindusiate, dorsal on veinlets, uniserial along each side of costule or midrib. Spores bean-shaped, perispore surface rugate, folds lobed, foveolate or sometimes reticulate. X=40.
Two sections, 10 species, 8 varieties and several hybrids in the world. Occurring in the temperate zone of north hemisphere (Asia, North America, and Europe) and the subtropical mountains of Asia, under forest. 5 species have been known from China.
Key to the species
1a. Laminae simple-pinnatifid, sori bigger, oblong. Most provinces and regions on the south of the Mt. Qinling
1. G. oyamense
1b. Laminae 2-pinnate to 3-pinnate, sori smaller, rotund.
2a. Rhaches more or less glandular abaxially.
3a. Raches sparsely glandular abaxially, densely glandular on the base of rachis and the connective with the petiolules of basal 1--3 pairs of pinnae; lateral veins often forked on segments. NE, N, NW, mountains of SW and Xizang
2. G. jessoense
3b. Fronds all over, especially on stipe top, raches and costae densely clothed with short glands on the abaxial surface, veins usually simple in segments. Xinjiang (Tian Shan)
3. G. robertianum
2b. Raches glabrous, eglandulose.
4a. Laminae nearly pentagonal-oval, ternate-formed, the lowest two pinnae almost same with the rest part of lamina in size, usually 2-pinnate-pinnatifid, the basiscopic basal pinnules of the lowest pinnae nearly same with the third pairs of pinnae in size, ultimate pinnules oblong, more or less pinnatifid, or波状 shallowly sinuate-lobed at margin. NE, NW and Nei Mongol
4. G. dryopteris
4b. Laminae deltoid-ovate, the lowest pinnae far smaller than the rest part of lamina, usually 3-pinnate-pinnatifid, the basiscopic basal pinnules of the lowest pinnae nearly same with the fourth pairs of pinnae in size, ultimate pinnules narrowly oblong, usually entire at margin. Taiwan and Yunnan
5. G. remotepinnatum
1. Gymnocarpium oyamense (Bak.) Ching in Contr. Biol. Lab. Sci. Soc. China Bot. Ser. 9:40, f. 3. 1933.
东亚羽节蕨 dong ya yu jie jue
Polypodium oyamense Bak. in Journ. Bot. 6:366. 1877; Dryopteris oyamensis C. Chr.; Currania oyamensis Copel.; Phegopteris oyamensis v. A. v. R.; Carpogymnia oyamensis Lφve et Lφve; Polypodium krameri Franch. et Sav.; Nephrodium krameri Diels; Phegopteris krameri Makino; Aspidium krameri Christ; Polypodium krameri var. incisum Franch. et Sav.; Polypodium gymnogrammoides Bak.; Nephrodium gymnogrammoides Diels; Dryopteris gymnogrammoides C. Chr.; Currania gracilipes Cop.; Gymnocarpium gracilipes (Cop.) Ching; Gymnocarpium oyamense var. gracilipes W. C. Shieh.
Rhizomes long-creeping, 1.5--2(--3) mm thick, clothed with red-brown broadly lanceolate scales, glabrate when old. Fronds distant; fertile fronds (13--)25--45(--50) cm; stipes (7--)12--25(--31) cm long, 1.5--2 mm thick, stramineous, lustrous, round abaxially and grooved adaxially, scaly at base, upward glabrate, articulate to lamina at top; laminae ovato-triangular, (7--)10--18(--22) cm long, (4--)6--13(--20) cm broad, acuminate at apex, cordate at base, pinnate parted to 44mm far from raches; segments 6--9(--13) pairs, opposite, patent, close each other, broadly falcate-lanceolate, curved up at apex, acute or acuminate at apex, decurrent to narrow costal wing at base, shallowly lobed to pinnatipartite at margin; the basal pair of segments often broadly lanceolate, descendant, curved at apex, (2--)4--7(--10) cm long, 1--1.8(--3) cm broad, the second pair of segments a little longer than the first pair, or nearly equal, patent, curved at apex too, from the third pairs segments upward gradually smaller, slightly upswept; secondary segments in some cases different in size, obtuse-rounded at apex, entire or crenate at margin. Veins thin in segments, pinnate, lateral veins (2--)4--5(--8) pairs, upswept, simple, slightly visible. Laminae herbaceous when dried, glabrous, green adaxially, pale-green abaxially; between rachis bases and stipe apices prominently articulate, stipes at upper part, raches at lower parte and the base of costules of lowest segments sparsely short-glandular pubescent, or absent. Sori oblong, medial on veinlets, biserial along two sides of costules, spaced each other. Spore surface rugate , foveolate. 2n=80,160.
Damp area under forests or in mosses on rocks; 300--2900 m. Shaanxi, Gansu, Anhui, Zhejiang, Jiangxi, Taiwan, Henan, Hubei, Hunan, Sichuan, Chongqing, Guizhou, Yunnan, Xizang [Nepal, Japan, Philippines, New Guinea]. Type from Japan.
2. Gymnocarpium jessoense (Koidz.) Koidz. in Acta Phytotax. Geobot. 5:40. 1936.
羽节蕨 yu jie jue
Dryopteris jessoensis Koidz. in Bot. Mag. Tokyo 38:104. 1924. (type KYO!); Dryopteris linnaeana var. jessoensis (Koidz.) C. Chr.; Lastrea jessoensis Akasawa; Carpogymnia jessoensis Lφve et Lφve; Aspidium dryopteris (L.) Baumg. var. longulum Christ; Gymnocarpium robertianum var. longulum (Christ) H. Ito ex Nakai; Gymnocarpium longulum Kitagawa; Lastrea robertiana var. longula (Christ) Ohwi; Gymnocarpium robertianum (Hoffm.) Newman subsp. longulum (Christ) Toyokuni; Dryopteris linnaeana C. Chr.; Dryopteris robertiana sensu C. Chr.; Gymnocarpium robertianum sensu Nakaike, pro parte; Polypodium dryopteris L. var. disjunctum Rupr.; Gymnocarpium dryopteris Newman var. disjunctum Ching; Gymnocarpium disjunctum Ching, pro parte; Gymnocarpium dryopteris subsp. disjunctum Sarvela; Carpogymnia disjuncta Lφve et Lφve; Gymnocarpium remotum Ching, pro parte; Gymnocarpium remote-pinnatum sensu Ching; Gymnocarpium continentale auct. non Pojark.
Rhizomes long-creeping, clothed with brownish ovato-lanceolate scales at apex; fronds usually distant, sometimes approximate. Fertile fronds (16--)20--50(--76) cm; stipes (8--)15--32(--51) cm long, up to 3.5 mm thick, stramineous, sparsely bearing scales at base, glabrate upwards; laminae deltoid-ovate, (7--)15--20(--27) cm long, (7--) 14--22(--30) cm broad, acuminate at apex, rounded at base, pinnate-pinnatifid or 2-pinnate-pinnatipartite; pinnae(3--)5--8 pairs, opposite, upswept, basal 1--4 pairs petioled, articulate to raches, basal pair of pinnae largest, narrowly triangular, (4--)8--15(--18) cm long, 3--7(--11) cm broad at base, acuminate at apex, near truncate at base, petiolules (0.8--)1--2.5(--3.5) cm, slightly upswept, pinnate-pinnatifid or pinnate-pinnatipartite; pinnules 5--8 pairs, deltoid-lanceolate, acuminate at apex, near truncate at base, opposite or subopposite, basal one to several pairs articulate to costae, usually sessile, in some cases the basal pair shortly petioled, 1--3(--12) mm, the basal basiscopic pinnules of pinnae the longest, 1--5(--7) cm long, (0.7--)1--2.3 cm broad, descendant, pinnate or pinnatifid; segments 5--10 pairs, oblong to narrowly ovate, rounded-obtuse at apex, bases free or joining by narrow wing each other, crenate at margin; the second pair of pinnae is (2--)4--5(--7.5) cm apart from the basal pair, narrowly triangular, far smaller than the lowest pinnae, 4--8(--12) cm; from the third pairs upward pinnae broadly lanceolate, gradually smaller. Veins pinnate in segments, veinlets usually forked, sometimes simple, very upswept, visible. Laminae herbaceous or papery when dried, pale-green; stipes top, raches, costae more or less bearing hyaline or pale yellow short glands. Sori small, round or oblong, dorsal on veinlets, close to the margin of ultimate pinnae or segments. Spore surface rugate, foveolate. 2n=80.
Wet areas under forests or mountain slopes; 400--4000 m. Heilongjiang, Jilin, Liaoning, Nei Mongol, Hebei, Shanxi, Henan, Shaanxi, Ningxia, Gansu, Qinghai, Xinjiang, Sichuan, Guizhou, NW Yunnan, SE Xizang [Afghanistan, N Pakistan, N India, Nepal, Sikkim, Korea, Japan, E Russia; NW North America]. Type from Japan.
3. Gymnocarpium robertianum (Hoffm.) Newman in Phytologist 4:371. 1851.
密腺羽节蕨 mi xian yu jie jue
Polypodium robertiana Hoffm. Deutsch. Fl. 2:20. 1795; Phegopteris robertiana Fee; Aspidium robertianum Luerss.; Dryopteris robertiana C. Chr.; Polypodium dryopteris var. robertianum Hook. et Bak.; Carpogymnia robertiana Lφve et Lφve; Polypodium dryopteris var. glandulosum Neilet; Polypodium calcareum J. Sm.; Aspidium calcareum Baumg.; Phygopteris calcarea Fee; Polypodium dryopteris var. calcareum Gray; Gymnocarpium altaycum C. Y. Yang.
Rhizomes long-creeping, clothed with brownish ovato-lanceolate scales at apex. Fronds distant; fertile fronds 18--40 cm; stipes 11--25 cm long, 0.5--1 mm thick, stramineous, sparsely bearing scales at base, glabrate upwards; laminae deltoid-ovate, 7--13 cm long, 4--8 cm broad, acuminate at apex, rounded at base, 2-pinnate to 2-pinnate, pinnules pinnatipartite or pinnatisect; pinnae 5--8 pairs, opposite, upswept, basal 3--4 pairs petioled, articulate to raches, the basal pair of pinnae the largest, narrowly triangular, 4.5--8.5 cm long, 3--4.5 cm broad at base, acuminate at apex, near truncate at base, petiolule 6--13 cm, slightly upswept, pinnate, pinnules pinnatipartite or pinnatisect; pinnules 4--6 pairs, deltoid-lanceolate or narrowly oblong, acuminate at apex, broadly cuneate at base, opposite or subopposite, basal1--2 pairs articulate to costae, subsessile, the basal basiscopic pinnules of pinnae the longest, 1.5--2.7 cm long, 6--8 mm broad, descendant, pinnate or pinnatipartite; segments 5--7 pairs, oblong to narrowly ovate, rounded-obtuse at apex, separate or connate with wings of costules at base, crenate at margin; the second pair of pinnae is 1.7--3.3 cm apart from the basal pair, narrowly triangular, far smaller than the basal pair of pinnae, 2--5.5 cm; pinnae from the third pairs upward broadly lanceolate, gradually smaller. Veins pinnate in segments, veinlets usually simple, sometimes forked, upswept, visible. Laminae herbaceous or papery when dried, pale-green adaxially, more pale-green abaxially; all over especially upper parte of stipes, lower parte of raches, costae and midribs shortly glandular pubescent. Sori small, rotund or oblong, dorsal on veinlets, submarginal. 2n=160.
Shad places under forests or mountain slopes; 1500--2500 m. Qinghai and Xinjiang [Wide spread in Europe and North America, Pakistan]. Type from Europe (Germany).
4. Gymnocarpium dryopteris (L.) Newman in Phytologist 4:371. 1851.
欧洲羽节蕨 ou zhou yu jie jue
Polypodium dryopteris L., Sp. Pl. 2:1093. 1753 (Neotype BM!); Lastrea dryopteris Bory; Aspidium dryopteris Bunmg.; Dryopteris dryopteris Christ; Carpogymnia dryopteris Lφve et Lφve; Dryopteris linnaeana C. Chr.
Rhizomes slender, creeping, black, lustrous, clothed with brown ovato-lanceolate scales at apex. Fronds distant; fertile lamina (15--)20--30(--50) cm; stipes 10--22(--35) cm, slender, stramineous, sparsely bearing scales at base; lamina pentagonal-oval or broadly ovato-triangular, broad and long nearly equal, 7--15(--20) cm, acuminate at apex, broadly cuneate at base, nearly tripartite, usually 2-pinnate-pinnatifid; basal pair of pinnae nearly the same size as the upper rest portion of lamina, narrowly triangular, (3.5--)5--9(--12) cm long, 2.5--4(--7) cm broad, pinnate-pinnatifid, subtruncate at base, with (0.8--)1--1.5(--2.5) cm stalks, the stalk of middle portion ca. 3 cm; pinnules 5--6 pairs, oblong-lanceolate, 1.5--2(--4) cm long, 0.5--2 cm broad, acute or acuminate at apex, rotund-cuneate at base, sessile, opposite or subopposite, patent; the largest pinnules with 6--10 pairs of segments, segments close each other, oblong to narrowly ovate, ca. 4 mm, rounded-obtuse at apex, joining by narrow wing each other at base, entire to shallowly lobed at margin; the second pair of pinnae is 1.5--4 cm apart from the basal pair, in some cases short petiole present, upward pinnae sessile. Veins pinnate in segments, veinlets simple, upswept, visible abaxially. Laminae thin-herbaceous or near membranous when dried, green, raches and costae slender, without glands. Sori small, exindusiate(1), rotund, dorsal on veinlets, uniserial along each side of midrib. Spore surface rugate, foveolate. 2n=160.
Damp areas under coniferous forests; 300--2900 m. Heilongjiang, Jilin, Liaoning, Nei Mongol, Shanxi, Shaanxi, Xinjiang [wide spread in the temperate zone of north hemisphere]. Type from Europe (Sweden).
Chromosome and allozyme evidences have shown that: Gymnocarpium dryopteris is an allotetraploid come from hybridization between Asian diploid G. jessoense and North American diploid G. appalachianum.
────────────────
(1) J. Sarval noted in 1976 that, indusia were found in a specimen of G. dryopteris from Japan, they were reniform, 0.5 mm in diameter, hyaline, glabrous, erose at margin. It may be atavism.
5. Gymnocarpium remotepinnatum (Hay.) Ching in Bull. Chin. Bot. Soc. 1(2):14. 1935.
细裂羽节蕨 xi lie yu jie jue
Dryopteris remote-pinnata Hay., Ic. Pl. Form. 6:Suppl. 108. 1916. (Type TI! topotype TAIF); Thelypteris remote-pinnata Alston; Dryopteris remota auct. non Hayek 1908; Gymnocarpium remotum Ching, pro parte.
Rhizomes slender, creeping, sparsely clothed with brownish ovato-lanceolate scales. Fronds distant; fertile fronds 20--30 cm; stipes (10--)12--20 cm, stramineous, slim and fragile, sparsely bearing scales at base; laminae triangular, 10--14 cm long, 6--9 cm broad, acuminate at apex, near truncate at base, usually 3-pinnate to 3-pinnate-pinnatifid; pinnae 5--6 pairs, opposite, the basal two pairs shortly petioled, the uppers sessile, the basal pair of pinnae the largest, narrowly triangular, 6--8 cm long, 5--6 cm broad, acuminate at apex, near truncate at base, petiolules 1--2.2 cm, bases articulated, patent, 2-pinnate; pinnules 5--6 pairs, opposite, oblong-lanceolate, 1.5--2 cm long, ca. 1 cm broad, acuminate at apex, broadly rounded at base, sessile, patent, pinnate; segments or secondary pinnules narrowly oblong, rounded-obtuse at apex, separate or connate with wings of costules at base, entire at margin, but cut at basiscopic side; the second pair of pinnae is 2--3 cm apart from the basal pair, oblong-falcate, smaller, ascendant upwards, rotund and symmetrical at base, sessile, upper pinnae in same shape with the second pair of pinnae but gradually smaller. Veins pinnate in segments, veinlets simple, very upswept, visible abaxially. Laminae thin-herbaceous when dried, greenish brown, all over glabrous, raches and costae on the abaxial surface without glands. Sori small, round, brown, exindusiate, dorsal on veinlets. 2n=80@.
* Under coniferous forests or on rocks in forest margin; 2500--3400 m. mountains Taiwan, NW Yunnan. Type from Taiwan.
5. ANISOCAMPIUM Presl, Epim. Bot. 52. 1849.
安蕨属 an jue shu
ATHYRIUM Kato et Kramer, pro parte.
Terrestrial medium- to small-sized plants. Rhizomes long-creeping or erect, clothed with brown lanceolate scales. Fronds distant or caespitose; stipes long, usually stramineous, sparsely bearing scales at base, glabrate upwards, grooved adaxially, open to raches; laminae ovato-oblong or deltoid-ovate, pinnate; pinnae 2--7 pairs, pinnatifid at apex or terminal pinna conform to pinnae in shape, the basals opposite or subopposite, the uppers alternate, falcate-lanceolate, acuminate at apex, symmetrical at base, the basals petioled, shallowly lobed at margin, segments toothed. Veins pinnate in segments, lateral veins 3--5 pairs, simple or occasionally forked, free, in some cases veinlets connivent at end as triangular areolae in lower 1--2 pairs. Laminae papery when dried, glabrous adaxially, on the abaxial surface costae or midribs bearing brown linear-lanceolate small scales and short pale hairs. Sori round, medial on veinlets, uniserial along each side of midrib or only 3--5; indusia small, round-reniform, membranous, ciliated at margin, fugacious. Spores bilateral, perispore surface with lophate ridges. X=40.
Three species known, in the tropics and subtropics of SE Asia; 2 species in China, the Yangtze River valley and the south of it.
Since the sori round, and veinlets in some cases connivent as triangular areolae in this genus, the systematic position of the genus has been changed for several times. However, morphological, cytological and molecular data show that this genus is most close to the genus ATHYRIUM Roth.
1a. Laminae pinnatifid at apex; veins free, veinlets terminating in the cartilaginous long spine-like teeth. the Yangtze River valley and S, SW China
1. A. sheareri
1b. Laminae imparipinnate, terminal pinna conform to other pinnae; the basal 1--2 pairs of veinlets in segments often connivent at tip forming triangular areolae, veinlets not terminating in teeth. Yunnan
2. A. cumingianum
1. Anisocampium sheareri (Bak.) Ching in Y. T. Hsieh in Acta Bot. Yunnan. 7(3):314. 1985.
华东安蕨 hua dong an jue
Nephrodium Sheareri Bak. in Journ. Bot. 200. 1875; Dryopteris Sheareri C. Chr.; Athyrium sheareri Ching; Nephrodium polypodiforme Makino; Dryopteris polypodiforme C. Chr.; Aspidium otarioides Christ; Dryopteris otarioides C. Chr.; Dryopteris subsagenioides Christ; Nephrodium isolatum Bak.
Rhizomes long-creeping, sparsely clothed with brownish lanceolate scales. Fronds approximate or distant; fronds 25--60 cm; stipes 15--30 cm long, ca. 2 mm thick at base, sparsely bearing scales, upward stramineous (occasionally pale red-purple), glabrate; laminae ovato-oblong or ovato-triangular, 15--30 cm long, 12--18 cm broad at middle, acuminate at apex, subtruncate or rotund-cuneate at base, pinnate, pinnatifid at apex; lateral pinnae 2--7 pairs, falcate-lanceolate, 6--10 cm long, 1.5--2 cm broad, long-acuminate at apex, rounded at base, the basal 1--2 pairs of pinnae often cuneate at the acroscopic bases, pinnatilobate to pinnatisect at margin, segments broadly ovate or oblong, with long-teeth at margin, upward segments gradually smaller. Veins free, pinnate in segments, lateral veinlets 3--4 pairs, simple or occasionally forked, extend into cartilaginous long teeth, only the basal pair of veinlets reach to the sinus or incision between teeth. Laminae papery when dried, glabrous adaxially, on the abaxial surface costae and costules bearing brownish small scales and pallid short hairs. Sori round, 3--4 pairs per segment, biserial along two sides of costules; indusia round-reniform, brown, membranous, ciliated at margin, fugacious. Spores perispore surface with lophate ridges. 2n=160,120 (3X apo.).
In valley under forests, streamside or shady mountain slopes; 1800--1900 m. Gansu, Jiangsu, Anhui, Zhejiang, Jiangxi, Fujian, Hubei, Hunan, Guangdong, Guangxi, Sichuan, Chongqing, Guizhou, Yunnan [Japan, South Korea]. Type from Jiangxi (Lu Shan).
2. Anisocampium cumingianum Presl, Epim. Bot. 59. 1849. (Isotype BM!)
安蕨 an jue
Athyrium cumingianum Ching; Aspidium otarium Kze et Mett.; Nephrodium otarium Bak.
Rhizomes short, erect, densely clothed with brown lanceolate scales at apex. Fronds caespitose; fronds 27--70 cm; stipes 12--45 cm long, 2--2.5 mm thick at base, densely clothed with scales, upward stramineous, glabrate; laminae broadly ovate or oblong, 15--25 cm long, ca. 10 cm broad at middle, acuminate at apex, subtruncate or rotund-cuneate at base, imparipinnate; terminal pinna conform to pinnae present or nearly imparipinnate, lateral pinnae 4--8 pairs, petioled, falcate-lanceolate, 6--10 cm long, 1.5--2 cm broad, acuminate at apex, rotund-cuneate at base, shallowly lobed at margin, near pinnatisect at lower margin, serrate at top margin; segments triangular, with several sharp teeth at apex. Veins free, pinnate in segments, lateral veins 4--5 pairs, simple, reaching into teeth, only the basal 1--2 pairs of veinlets reaching to underneath sinus, and often connivent at tip forming triangular areolae. Laminae thin papery when dried, glabrous adaxially, abaxial costae and midribs sparsely bearing brown small scales and pale short glandular-hairs. Sori round, 3--4(--5) pairs per segment, biserial along two sides of costules, a little close to midrib; indusia round-reniform, brown, membranous, ciliated at margin, fugacious. Spore perispore surface with lophate ridges, sometimes connected into reticulate or ornate. 2n=80,160.
Damp areas under evergreen forests, on limestone rocks; 700--1000 m. S Yunnan, Taiwan [Laos, S India, S Myanmar, Philippines, Indonesia (Java) and Sri Lanka]. Type from Philippines (Luson).
6. KUNIWATSUKIA Pic. Ser. in Webbia 28:455. 1973.
拟鳞毛蕨属 ni lin mao jue shu
MICROCHLAENA Ching; ATHYRIUM Kato et Kramer, pro parte.
Terrestrial medium-sized herbal plants. Rhizomes short, erect or somewhat ascending, lignous, densely clothed with brown linear scales at apex. fronds caespitose; fronds radial arrangement, patent. Stipes long, pale purple when young, thick at base, section triangular, 2 vascular bundles present, upward conjoint as U-shaped; laminae oblong, pinnate, subimparipinnate at tip, or imparipinnate (terminal pinna conform to lateral pinnae, petioled), lateral pinnae ca. 20 pairs, alternate, ascendant, linear-lanceolate, shallowly lobed or incised-serrulate at margin. Veins free, visible on surfaces, lateral veins pinnate, veinlets 3--6 pairs, simple, catadromic, except the basal basiscopic veinlets come out directly from costule, not extend to the sinus between teeth, the acroscopic veinlets come from the base of lateral veins, reach to the half way and stop. Laminae papery when dried, glabrous on surfaces, costae grooved adaxially, protuberant abaxially, occasionally with a few brown small scales. Sori small, round, dorsal on the base or the lower part of veinlets; indusia small, round-reniform, thin-membranous, lacerate as ciliate at margin, fugacious. Spores bilateral, perispore surface with lophate ridges. X=40.
Since its sori round, this genus was often been put in family DRYOPTERIDACEAE systematically before. In fact, its most characters and basal chromosome number are the same to family ATHYRIACEAE, and it is very close to the genus ANISOCAMPIUM Presl.
Only one species. SW China [N India, N Myanmar, Nepal, Bhutan, Sikkim, Thailand, Sri Lanka].
1. Kuniwatsukia cuspidata (Mett.) Pic. Ser. in Webbia 28:455. 1973.
拟鳞毛蕨 ni lin mao jue
Aspidium cuspidatum Mett. in Abhandl. Senckenb. Naturf. Ges. (Frankfurt) 2(2):Aug. 1858, non Desv., nom. nud. 1827; Nephrodium cuspidatum Bak.; Lastrea cuspidata Bedd.; Dryopteris cuspidata Christ; Microchlaena cuspidata Ching; Athyrium cuspidatum Kato; Polypodium elongatum Wall. ex Hook.; Phegopteris elongata J. Sm.; Lastrea elongata Bedd. ex Clarke; Dryopteris elongata O. Ktze.; Polypodium trinidadensis Jenm.; Dryopteris trinidadensis C. Chr.; Aspidium yunnanense Christ; Microchlaena yunnanensis Ching; Dryopteris yunnanensis Cop.; Dryopteris khasiana C. Chr.
Rhizomes short, erect or somewhat ascending, densely clothed with brown linear scales at apex. Fronds caespitose; fronds up to 1 m; stipes 30--65 cm long, up to 5 mm thick at base, sparsely bearing scales, pale purple or brownish upward, glabrate; laminae oblong-lanceolate, 35--70 cm long, 16--30 cm broad, acuminate at apex, rotund-cuneate at base, imparipinnate, lateral pinnae 7--23 pairs, alternate, petioled, ascendant, lanceolate, 12--18 cm long, 1.5--2 cm broad, long-acuminate at apex, rotund-cuneate or asymmetrically cuneate at base, incised-serrulate or serrate at margin, tooth tip beak-like and with 1 or 2 small teeth; terminal pinna conform to pinnae present, a little bigger, usually there are 1 or 2 small segments at base, the lower part shallowly lobed at margin, segments obtuse-rounded, with 1--3 small teeth at apex. Veins free, visible on surfaces, lateral veins pinnate, veinlets 3--6 pairs, simple, catadromic, except the basal basiscopic veinlet come out directly from midrib, not extend to the sinus between teeth, the acroscopic veinlet come from the base of lateral veins, reach to the half way and stop. Laminae papery then dry, dark green, upper surface glabrous, costae occasionally bearing a few brown small scales abaxially. Sori small, dense, round, cinnamon, dorsal on the base or lower part (occasionally on the middle) of veinlet; indusia small, round-reniform, brown, thin-membranous, lacerate as ciliate at margin, fugacious. Spore perispore with lophate ridges. 2n=80.
Damp area under evergreen broad leaf forests or shrubs; 500--1800(--2000) m. Guangxi, Guizhou, SE Yunnan, SE Xizang [Nepal, Bhutan, Sikkim, N India, N Myanmar, N Thailand, Sri Lanka, W Himalayas].
Hooker & Bak. (1868) thought that the type specimen (K!) collected from Kew Garden originally was come from Sri Lanka, but Clarke (1880) didnt think so.
7. PSEUDOCYSTOPTERIS Ching in Acta Phytotax. Sin. 9:76. 1964.
假冷蕨属 jia leng jue shu
CYSTOPTERIS Maxim. pro parte; ATHYRIUM Milde. pro parte.
Rhizomes long-creeping, forked-branched, blackish brown, bearing remainder stipe bases, sparsely scaly, more dense scaly at apex, scales ovate to ovato-lanceolate brownish. Fronds separate; stipes long, rarely short, sparsely scaly, black, a little attenuate, but not swollen as spindle at base; laminae broadly ovato-triangular, or oblong-ovate, rarely oblanceolate, 3-pinnate to 4-pinnate, rarely pinnate to 2-pinnate-pinnatifid, pinnatifid-acuminate at apex; pinnae lanceolate, broadly oblanceolate, rarely ovate or elliptic, the basal pair of pinnae not shortened or only a little shortened, more or less narrowed towards pinna base, shortly petiolulate, in a few cases lower pinnae gradually shortened, not narrowed toward the pinna base, subsessile, ultimate segments serrate; veins pinnate at segments, lateral veins free, usually forked, upward simple, 1 veinlet per tooth; laminae herbaceous or hard herbaceous when dried, raches bearing short glandular-hairs or sub-glabrous, rarely with sparsely long multicellular hairs abaxially; costules and midribs without spine-like processes on the adaxial surface. Sori small, oblong or rotund, sub-lateral or dorsal on veinlets, often lateral on veinlets at top of pinnules; indusia round-reniform or oblong, laciniate or ciliated at margin. Spores bilateral, perispore surface with folds. X=40.
About 7 species in the world, all in China. Mainly occur in China, in the temperate zone, subtropic mountains and the Qinghai-Xizang Plateau [N India, Korea, Japan and E Russia].
This genus was set up by Renchang Ching in 1964 based on the species Cystopteris spinulosa Maxim from North-eastern Asia. However, most species of the genus is confined in West China and the Himalayas. It is morphologically between the genera CYSTOPTERIS Bernh. and ATHYRIUM Roth. It is different form the former in muck taller, ampler leaves of deltoid-ovate outline with sharply serrated leaf-margin, round-reniform or reniform indusia with broad subcordate base and ciliated margin, spores not echinate but winged on the surface, and basal chromosome number of 40 but not 42; different from the later in rhizome long-creeping with a few scales, stipe not swollen as spindle and beak-shaped at base, sori nearly round, dorsal on veinlets, rarely sub-lateral on veinlets, indusia round-reniform or reniform. This genus are overlapped with the later in some of characters, and shared the same basal chromosome number 40, in addition, molecular data also shows that it is very close to the genus ATHYRIUM Roth.
1a. Large- to medium-sized plants, leaf length usually more than 60--70 cm, up to 2--3 m, all pinnae petiolulate. Mountains in the south of Mt. Qinling.
1. P. atkinsonii
1b. Medium- to small-sized plants, leaf length usually less than 60--70 cm, from the second pair of pinnae upward all pinnae subsessile or very shortly petiolulate. Mountains from NE China to SE Xizang.
2a. Laminae triangular, the basal pair of pinnae the largest, long-elliptic to elliptic-oblanceolate, their lower pinnules gradually shortened towards the base, prominently narrowed at base, pinnae above the second pair their pinnules often catadromic. Mostly occur in the mountains below 3000 m alt. 3a. Raches and costae glabrous abaxially, pinnules roundish at base, distant from costae, teeth long and pointed up at segment margin. NE, Nei Mongol and Mt. Qinling.
2. P. spinulosa
3b. Raches and costae more or less glandular-hairy abaxially, pinnules subtruncate at base, often close to costae, teeth short and stretched at segment margin. Shaanxi and S Gansu W Hubei, Sichuan, Yunnan and SE Xizang
3. P. subtriangularis
2b. Laminae ovate to oblanceolate, basal pair of pinnae more or less shortened, ovate to oblong-lanceolate, rarely near auriform, their lower pinnules not gradually shortened toward base, at most 1--2 pairs a little shortened, pinnules or segments of all pinnae anadromic or subopposite. Only occur in mountains above 3000 m alt.
4a. Laminae ovate to ovato-oblong, usually 2-pinnate to 3-pinnate; pinnule asymmetrical at base, shortly petiolulate; raches and costae bearing no long multicellular hairs.
5a. Fronds less than 35 cm; ultimate segments more close each other, teeth short at margin. W Sichuan, NW Yunnan and SE Xizang
4. P. schizochlamys
5a. Fronds up to 55 cm; ultimate segments thinner, separate each other, teeth long and thin at margin. NW Yunnan
5. P. atunzeensis
4b. Laminae oblanceolate or ovato-oblong, pinnate-pinnatifid to 2-pinnate, segments joining each other by narrow wing at base, if bippinate, pinnules nearly symmetrical at base, sessile; raches, costae and veins with sparse long multicellular hairs.
6a. Laminae oblanceolate, many pairs of lower pinnae gradually shortened towards the base. W Sichuan, NW Yunnan and SE Xizang
6. P. davidii
6b. Laminae ovato-oblong, only the basal 2 pairs of pinnae a little shortened. Sichuan (Lushan)
7. P. repens
1. Pseudocystopteris atkinsonii (Bedd.) Ching in Acta Phytotax. Sin. 9(1):78. 1964.
大叶假冷蕨 da ye jia leng jue
Athyrium atkinsonii Bedd. in Ferns Brit. S. Ind. Suppl. II, t. 359. 1876 (Type K!) & Handb. Ferns Brit. Ind. 163. 1883; Asplenium atkinsonii Clarke; Polypodium foliolosum Wall., nom. nud. (Syntype K!); Asplenium andersonii Clarke; Asplenium atkinsonii Bedd. var. andersonii Clarke; Pseudocystopteris andersonii Ching; Athyrium andersonii Panigr. et Basu; Asplenium fimbriatum Wall. var. sphaeropteroides Clarke; Athyrium fimbriatum var. sphaeropteroides Bedd.; Aspidium senanense Franch. et Sav.; Dryopteris senanensis C. Chr.; Athyrium senanense Koidz. et Tagawa; Asplenium lastreoides Bak.; Athyrium lastreoides Diels; Davallia athyrifolia Bak.; Polypodium leptophyllum Bak.; Dryopteris gracilifrons C. Chr.; Leucostegia yaklaensis Bedd.; Davallia yaklaensis C. Chr.; Araiostegia yaklaensis Nayar et Kaur; Athyrium microsorum Makino; Athyrium filix-femina Roth var. deltoidea Christ; Aspidium fargesii Christ; Dryopteris fargesii C. Chr.; Athyrium monticola Rosenst; Cystopteris grandis C. Chr.
Rhizomes short-creeping, 0.5--1 cm thick, bearing remainder stipe bases, young fronds clothed with more brownish or brown ovate scales at apices and stipe bases. Fronds approximate or distant; fertile fronds (15--)60--100(--200) cm; stipes 20--50(--100) cm long, 3--10 mm thick, blackish brown at base, upward gradually changed to stramineous or pale red-purplish, occasionally with thin scales; laminae usually larger, (24--)35--50(--100) cm, nearly as broad as long or a little narrowed, broadly ovate or ovato-triangular, in small plants laminae often ovate, while in large plants laminae often subtriangular, acuminate at apex, mature lamina 2-pinnate to 4-pinnate, raches and costae slightly tortuous; pinnae 10--12(--20) pairs, alternate, upswept, petiole 0.5--1(--3) cm, separate each other, somewhat as imbricate, the basal pair of pinnae is similar to the second pair, 5--15 cm apart from each other, 15--20(--50) cm long, up to 6--8(--28) cm broad at middle, slightly narrowed toward base, narrowly ovate to long ovato-lanceolate, acuminate at apex, near truncate at base, 2-pinnate-pinnatifid to 3-pinnate; pinnules 10 pair or more, anadromic, alternate, with ca. 2 mm short petiolule, 1.4--2 cm apart from each other, sub-patent, the basal pair a little shortened, the middles 3.5--5 cm long, 1.2--2 cm broad, oblong-lanceolate, acuminate at apex, truncate at base, pinnate-pinnatifid to 2-pinnate; secondary pinnules 6--8 pairs, 5--8 mm long, 2--3 mm broad, oblong or near ovate, obtuse-rounded and with unequal short teeth at apex, broadly cuneate and the basiscopic side slightly decurrent at base, pinnate or pinnatifid; ultimate segments ovato-oblong, rounded-obtuse or near square and with 23 unequal short teeth at apex, broadly cuneate and slightly asymmetrical at base. Veins visible on surfaces, more prominent abaxially, pinnate with 4--5 pairs of lateral veins on ultimate pinnules, veinlets simple, rarely forked. Laminae herbaceous when dried, blackish brown or greenish brown, glabrous on surfaces, the groove of costule is open to the groove of midribs adaxially, raches and costae occasionally with membranous, small-lanceolate scales abaxially. Sori round or elliptic, dorsal or semi-lateral on the basal acroscopic veinlets of segments, usually only 1 veinlet per segment, 2--3 pairs per ultimate pinnule; indusia small, round-reniform, occasionally oblong or J-shaped, membranous, pale-brown, slightly erose at margin, fugacious, look like exindusiate when mature. Perispore surface with folds, concave-convex, ornate. 2n=80.
Damp areas under coniferous forests or shrubs; 1200--4000 m. S Shanxi, Shaanxi, Gansu, Jiangxi, Fujian, Taiwan, Henan, Hubei, Hunan, Chongqing, Sichuan, Guizhou, Yunnan, SE Xizang [Japan, S Korea, N Myanmar, Sikkim, Bhutan, Nepal, N India, N Pakistan, Kashmir, Himalayas]. Type from Sikkim.
2. Pseudocystopteris spinulosa (Maxim.) Ching in Acta Phytotax. Sin. 9:78. 1964.
假冷蕨 jia leng jue
Cystopteris spinulosa Maxim. in Mem. Acad. St.-Petersb. 9:340. 1859 (Type K!); Athyrium spinulosum Milde; Asplenium spinulosum Bak.
Rhizomes long-creeping, ca. 2.5 mm thick, blackish brown, sparsely clothed with broadly ovate membranous scales. Fronds distant; fertile fronds 45--50 cm; stipes 22--26 cm long, 2--3 mm thick at base, same color with rhizomes, clothed with broadly ovate to lanceolate scales, often densely clothed with yellow long pubescence near the surface of soil, upward stramineous, occasionally with scales, glabrate, polished; laminae ovato-triangular to ovato-triangular, 20--28 cm, nearly as broad as long or a little broader, short-acuminate at apex or near acute, 2-pinnate to 3-pinnate; pinnae ca. 12 pairs, the basal pair the longest, 17--20 cm long, 6--8 cm broad at middle, elliptic to elliptic-oblanceolate, acuminate at apex, conspicuously gradually narrowed toward base, nearly symmetrical at base, with 5--8 mm short petiolule, subopposite, 2--3.5 cm apart from the upper pair, ascendant, pinnate to 2-pinnate; the basal pair of pinnae with anadromic pinnules 12--14 pairs, ascendant or explanate, oblong-lanceolate to ovato-triangular; the pinnae below lamina middle with gradually shortened pinnules toward base, obtuse or acuminate at apex, with long upward pointed teeth at margin, symmetrical, rounded or rotund-cuneate at base, sessile or slightly connate with costae, rarely very shortly petiolulate, the middle pinnules the longest, 4--5 cm long, 8--15 mm broad, pinnate; secondary pinnules no more than 10 pairs, upswept, oblong or narrowly ovate, 5--8 mm long, 2.5--3.5 mm broad, usually decurrent and connate with wings of costules at base, cut at margin, or with thin-long, sharp teeth at margin and apex, segments with the same teeth at margin; from the second pair upward pinnae lanceolate, narrower, subopposite or alternate, not narrowed toward base, upward gradually sessile; first pair of pinnules catadromic or subopposite, 2-pinnate; ultimate pinnules with the same pointed up, long, sharp teeth at margin. Veins visible on surfaces, pinnate in ultimate pinnules, veinlets simple, terminating into the tooth tip. Laminae herbaceous when dried, green or greenish brown, costules and midrib slightly with short spine-like processes on the adaxial surface, raches and costae sparsely bearing short hairs adaxially, glabrate abaxially. Sori round, dorsal on veinlets, 1 to several pairs on every ultimate segment; indusia near round-reniform, membranous, brownish. spore perispore thin, hyaline, surface with more folds. 2n=80.
Damp areas under coniferous, mixed forests or in shrubs, bamboo groves; 800--3000 m. Heilongjiang, Jilin, Liaoning, Nei Mongol, Shaanxi, Henan, Sichuan [Korea, Japan, E Russia]. Type from E Russia (Amur).
3. Pseudocystopteris subtriangularis (Hook. ex Hook. et Bak.) Ching in Acta Phytotax. Sin. 9:78. 1964.
三角叶假冷蕨 san jiao ye jia leng jue
Asplenium subtriangulare Hook. in Hook. & Bak., Syn. Fil. 225. 1867 (Type K!); Athyrium subtriangulare Bedd.; Asplenium spinulosum var. subtriangulare Clarke; Athyrium spinulosum var. subtriangulare C. Chr.; Asplenium spinulosum Clarke; Athyrium spinulosum sensu Bedd.; Athyrium hookerianum Moore; Athyrium longipes Christ; Pseudocystopteris longipes (Christ) Ching; Athyrium subtriangulare Bedd. var. sikkimensis Bir; Athyrium sikkimense Lφve et Lφve; Pseudocystopteris tibetica Ching; Pseudocystopteris remota Ching; Pseudocystopteris spinulosa var. taipaishanensis Ching; Pseudocystopteris sparsa Ching et S. K. Wu; Pseudocystopteris decipiens Ching et S. K. Wu; Pseudocystopteris reflexipinnula Ching et S. K. Wu; Pseudocystopteris purpurascens Ching et S. K. Wu.
Rhizomes long-creeping, 3--4 mm thick, blackish brown, sparsely scaly; scales brownish ovato-lanceolate. Fronds distant; fertile fronds (13--)60--70(--90) cm; stipes (5--)30--40(--55) cm long, (1--) 3--4(--5) mm thick at base, same coloured with rhizomes, densely scaly, often densely clothed with long yellow pubescence near surface of soil, upward stramineous, occasionally with brownish lanceolate membranous scales; laminae broadly ovato-triangular to ovato-triangular, (8--)25--35(--40) cm long, (10--)20--30(--45) cm broad at base, acuminate at apex, 2-pinnate to 3-pinnate; pinnae(10--)15--20 pairs, nearly alternate, upswept, close each other, basal pair of pinnae a little longer, more broad, 14--20(--28) cm long, 5--9(--12) cm broad at middle, elliptic to elliptic-oblanceolate, acuminate at apex, gradually narrowed toward base, nearly symmetrical at base, petiolules 5--8 mm, subopposite, (1--)4--5 cm apart from the upper pair, 2-pinnate; pinnules of the basal pair of pinnae anadromic, (10--)14--20 pairs, patent or slightly ascendant, with pinnules from pinna middle gradually shortened toward base, oblong- lanceolate or ovato-triangular, obtuse at apex, usually with patulous teeth; middle pinnae with the pinnules oblong-lanceolate, 2.5--4.5(--7) cm long, 1--1.5 cm broad at base, short-acuminate or acute at apex, sessile, subtruncate, separate or partly connate with costae, close each other, pinnate; secondary pinnules 10--12 pairs, patent or slightly upswept, oblong, 5--8 mm long, 2.5--4 mm broad, obtuse at apex, connate with wings of costules at base, cut at margin or with patulous teeth at apex and margin; the second pair of pinnae similar to but a little smaller than the basal pair, oblong; upward pinnae lanceolate, acuminate at apex, not narrowed toward base, pinnate-pinnatifid; ultimate segments serrate at apex and margin. Veins visible on surfaces, pinnate in ultimate pinnules, veinlets simple, terminating into tooth apices. Laminae herbaceous when dried, green or brown-green, costules and midribs on the adaxial surface with very short spine-like processes, raches, costae and costules more or less with short hairs, veins sparsely bearing glad-like short hairs or multicellular hairs abaxially, costae occasionally with a few small scales at base. Sori round, dorsal on veinlets, 1 to several pairs per ultimate segment; indusia membranous, round-reniform, brownish. Spore perispore thin, hyaline, with folds on surface. 2n=80@,160.
Damp area under forests; (500--)2000--4100 m. Shaanxi, Gansu, Qinghai, Hubei, Chongqing, Sichuan, Yunnan, SE Xizang [Nepal, Bhutan, Sikkim, N India]. Type from Sikkim.
4. Pseudocystopteris schizochlamys Ching in Acta Phytotax. Sin. 9:79. 1964 (Type E!).
睫毛盖假冷蕨 jie mao gai jia leng jue
Athyrium schizochlamys K. Iwats.; Pseudocystopteris sinica Ching; Pseudocystopteris lanpingensis Ching; Pseudocystopteris laterepens Ching.
Rhizomes long-creeping, 2--3 mm thick, densely clothed with brown ovato-lanceolate membranous scales at apex. Fronds distant; fertile fronds (9--)20--25(--40) cm; stipes (4--)6--10(--23) cm long, 1--3 mm thick, sparsely bearing scales at base, stramineous on the lower part, polished, often pale red-purplish on the upper part; laminae ovate to ovato-oblong, (5--) 11--17(--25) cm long, (3--)5--9(--20) cm broad, acuminate at apex, pinnate to 3-pinnate, pinnae or ultimate pinnules pinnatipartite; pinnae 10--13(--16) pairs, alternate, in some cases lower pinnae subopposite, the basal 1--2 pairs a little shortened, petioled, petiole 1--2(--4) mm, ascendant, curved upward in the top, (1--)1.5--2(--4) cm apart from each other, middle pinnae (1.5--)3--6(--11) cm long, (0.8--)1.5--2.4(--4) cm broad, oblong-lanceolate, nearly symmetrical at base, pinnate-pinnatifid; pinnules no more than 6--10 pairs, broadly ovate to ovato-lanceolate, close each other, basal pair of pinnae 8--10 mm long, 5--8 mm broad, with sharp teeth at apex, asymmetrical at base, the acroscopic side parallel to costae, the basiscopic side narrowly cuneate, slightly decurrent and connate with wings of costae, all anadromic, the basal 1--2 pairs pinnules of the lowest pinnae in some cases a little shortened; secondary pinnules or segments 3--4(--8) pairs, elliptic to oblong, 3--4(--7) mm long, ca. 2 mm broad, oblong, toothed at apex, cut at margin; ultimate segments with 23 teeth at apex; top pinnae oblong-lanceolate, obtuse and with sharp teeth at apex, slightly decurrent at base, 2-pinnate, pinnules shallowly lobed; ultimate segments toothed at apex. Veins visible abaxially, pinnate in pinnules, veinlets terminating in entire teeth. Laminae herbaceous or near papery when dried, dark-green, raches and costae usually pale red-purplish, glabrous or shortly glandular-hairy. Sori small, round or round-reniform, 1-3 sori per ultimate pinnule, the basal ultimate pinnules bigger, often 2 pairs present, dorsal on or semi-lateral on veinlets, on the basal acroscopic side of segments close to sinus; indusia round-reniform, brownish, lacerate as ciliated at margin. Spore perispore thin and hyaline, surface with a few folds. 2n=ca.80.
Damp areas under forests; 3000--4500 m. Sichuan, Yunnan, SE Xizang [Nepal]. Type from SE Xizang.
5. Pseudocystopteris atuntzeensis Ching, in Acta Phytotax. Sin. 9:83. 1964 (Type PE!).
阿墩子假冷蕨 a dun zi jia leng jue
Rhizomes long-creeping, ca. 2 mm thick, blackish brown, sparsely scaly, more densely scaly at apex and stipe bases, scales brown, broadly lanceolate. Fronds distant; fertile fronds 30--55 cm; stipes 12--23 cm long, 1--2 mm broad, stramineous, clothed with very sparse lanceolate small scales; laminae 15--32 cm, nearly as broad as long or a little narrowed, ovato-oblong or broadly ovato-triangular, acuminate at apex, pinnate shortly caudate at apex, 3-pinnate-pinnatifid; pinnae 8--10 pairs, ascendant, the lowest pinnae 10--16 cm long, 3--5 cm broad at middle, oblong-lanceolate, acuminate at apex, rotund-cuneate at base, anadromic, the acroscopic side often overlapping raches, the basiscopic side oblique, petiolules 1.5--3 mm, nearly opposite or alternate, 2.5--5 cm apart from each other, 2-pinnate-pinnatifid; pinnules ca. 9 pairs, shortly petiolulate, alternate, subpatent, spaced each other, the basal 1--2 pairs often a little shortened, ovato-triangular, the middles 1.5--3 cm long, 7--12 mm broad, oblong, with 2--3 thin-long teeth at apex, broadly cuneate, nearly symmetrical, or asymmetrical the acroscopic side parallel to costae at base, pinnate-pinnatifid; secondary pinnules ca. 5 pairs, oblique, distant, 4--7 mm long, 2--3 mm broad, oblong or rhomboid, with 23 long tooth at apex, broadly cuneate decurrent and connate with wings of costule at base, cut at margin; segments then and long, with 2--3 long tooth only at apex; pinnae gradually shortened upwards from middle of lamina, not narrowed, but often a little widened at base; upper pinnae oblong, sessile, pinnatifid, segments with several long tooth at apex. Veins visible on surfaces, pinnate in ultimate pinnules, veinlets terminating in entire teeth. Laminae herbaceous when dried, pale-green and glabrous; raches and costae pale red-purplish adaxially. Sori small, round, dorsal on suprabasal veinlets of segments, near the sinus between segments; indusia near round-reniform, pale-green, lacerate as ciliated-like at margin.
* On grassy slope; 3600--3700 m. NW Yunnan (Deqen).
6. Pseudocystopteris davidii (Franch.) Z. R. Wang in W. T. Wang (Ed.), Vasc. Pl. Hengduan Mount. 1:77. 1993.
大卫假冷蕨 da wei jia leng jue
Polypodium davidii Franch. in Nouv. Arch. Mus. Hist. Nat. Paris ser. 2. 10:119. 1887. (Type P!); Phegopteris davidii Bedd.; Athyrium davidii Christ; Polypodium dentigerum Wall. pro parte; Asplenium Duthiei Bedd.; Athyrium Duthiei (Bedd.) Bedd.
Rhizomes long-creeping, with remainder stipe bases, chestnut-black, glabrate, slightly polished, 1--2 mm thick, sparsely bearing red-brown, ovato-lanceolate scales at apices and stipe bases. Fronds approximate or distant; fertile fronds 20--35 cm; stipes 2.5--12 cm long, 1--2 mm thick, dark brown at base, upward stramineous, slightly pale red-purplish, clothed with a few scales; laminae oblanceolate or narrowly oblanceolate, 14--25 cm long, 3.5--8 cm broad, narrowed toward base, usually only 1--2(--3) cm broad, the widest at the upper-middle, acuminate at apex, pinnate; pinnae 13--24 pairs, sessile or subsessile, many pairs of lower pinnae gradually shortened, explanate or slightly weeping, subopposite, the basal pair of pinnae somewhat auricled, usually only 0.3--1.3 cm, the middle and upper pinnae slightly upswept, alternate, costae spaced in 0.5--3 cm from each other, the middle pinnae the largest, 1.5--4 cm long, 5--14 cm broad, lanceolate or broadly lanceolate, short-acuminate at apex, subtruncate, nearly lateral symmetrical at base, or the acroscopic segments only a little bigger, pinnatipartite to pinnatisect; pinnae no more than 10 pairs, ovate to ovato-triangular, 2--8 mm long, 2--5 mm broad, obtuse at apex, joining each other by narrow wing or slightly asymmetrical at base, with sharp and stretched tooth or shallowly lobed at margin, segments with 2--3 short tooth. Veins visible abaxially, red-purplish, lateral veins simple or 2--3-forked, extend to the tooth tip. Laminae herbaceous or near papery when dried, dark greenish-brown adaxially, pale-green abaxially. Raches and costae bearing a few long scale-like hairs abaxially, costae and veins bearing sparse long multicellular hairs on surfaces, costae more or less with short glandular-hairs abaxially. Sori elliptic or reniform, 1-7 sori per segment, setting on the bases of acroscopic veinlets of forked veins or the medium of simple veins; indusia small, thin-membranous, round-reniform or narrowly ovate, lacerate at margin, hidden by sporangia at maturity, hard seen. Spores bilateral, perispore surface with folds. 2n=80.
Alpine cool mountains, grassy slopes, under shrub groves or in rock slits; 3000--4300 m. W Sichuan, NW Yunnan, SE Xizang [N Myanmar, Nepal, Sikkim, N India, Himalayas]. Type from Sichuan (Baoxing).
This species originally was been put in genus ATHYRIUM. But it morphologically possesses the characters such as rhizomes blackish brown, almost bearing no scales, thin and long-creeping, stipes not swollen as spindle and beak-shaped at base, sori cystopterioid etc., and lives in the high altitude area, SichuanHimalayas, those are in keeping with the genus PSEUDOCYSTOPTERIS. Except its basal pinnae gradually shortened, all other characters are almost the same as Pseudocystopteris repens. In addition, there are some species with the character of the basal 1--2 pairs of pinnae a little shortened in the genus PSEUDOCYSTOPTERIS, such as P. schizochlamys and P. repens, especially bearing the special long multicellular hairs on the adaxial surface of raches, costae and veins in common, sympatric growing, which shows they have very close relationship. However, we can hard to find a species in the genus Athyrium so close to Athyrium davidii. That is why it was recombined to the genus PSEUDOCYSTOPTERIS.
7. Pseudocystopteris repens Ching in Acta Phytotax. Sin. 9:80. 1964 (Type PE!).
长根假冷蕨 chang gen jia leng jue
Rhizomes long-creeping, 2.5--3 mm thick, blackish brown, almost bearing no scales, only clothed with brownish, membranous, broadly ovato-lanceolate scales at apices and stipe bases. Fronds distant; fertile fronds 20--30 cm; stipes 7--10 cm long, 1.5--2 mm thick at base, same colour with rhizomes at base, upward gradually become red-purplish, sparsely bearing broadly lanceolate scales; laminae 14--17 cm long, 7--10 cm broad at middle, ovato-oblong, acuminate at apex, near truncate and slightly narrowed at base, 2-pinnate; pinnae 13--15 pairs, the lowest two pair a little shortened, slightly pendulous, 2.3--1.5 cm apart from each other, middle pinnae 4--5 cm long, 1.5 cm broad at base, broadly lanceolate, short-acuminate at apex, truncate at base, sessile, pinnate; pinnules 10--12 pairs, patent, close each other, ca. 7 mm long, ca. 4 mm broad, oblong-ovate, obtuse with triangular short teeth at apex, subsymmetrical and broadly cuneate at base, the basiscopic side decurrent and connate with narrow wings of costae. Duplicato-serrate, or cut at margin, segments with 2--3 short teeth; top pinnae narrowly oblong-lanceolate, joining each other by narrow wing at base, pinnatipartite, segments with short-triangular teeth at margin. Veins visible abaxially, pinnate in pinnules, 1 veinlet per tooth. Laminae herbaceous when dried, green, along raches and costae bearing sparse long multicellular hairs and short glandular-hairs on surfaces. Sori small, elliptic or round-reniform, dorsal on veinlets, 1-6 every ultimate pinnule or segment, medial; indusia round-reniform or narrowly ovate, brownish, lacerate at margin.
* Under azalea forests; 3200--3700 m. Sichuan. Type from Sichuan (Lu Shan).
8. NEOATHYRIUM Ching et Z. R. Wang in Acta Phytotax. Sin. 20(1):76. 1982.
新蹄盖蕨属 xin ti gai jue shu
CORNOPTERIS Kato et Kramer pro parte.
Medium-sized terrestrial. Rhizomes moderately stout, creeping, sparsely clothed with big brownish scales at apex and stipe bases. Fronds distant; stipes almost as long as lamina, moderately stout, not beak-shaped at base; laminae deltoid-ovate, acuminate at apex, slightly cordate at base, tripinnatipartite; pinnae lanceolate or oblong-lanceolate, lower pinnae shortly petiolulate, subopposite, ascendant, the basal 2 pairs of pinnae largest, acuminate at apex, a little narrowed toward base, with numerous pairs of pinnules; pinnules lanceolate, acuminate at apex, broadly cuneate at base, subsessile, the lowers subopposite, upwards alternate, patent or slightly ascendant; ultimate segments oblong, obtuse-rounded at apex, joining each other by narrow wing at base, serrulate at margin. Veins unclear adaxially in segments, but visible abaxially, pinnate, midrib slightly tortuous, lateral veins simple or forked. Laminae papery when dried, greenish-brown adaxially, green abaxially; fleshy hornlike process absent in adaxial grooves at the base of costules; raches, costae and costules bearing pallid unicellular short hairs and pale-brown multicellular long hairs on the abaxial surface. Sori round or elliptic, exindusiate, dorsal on the middle of veinlets. Spores bilateral, perispore surface with rugate processes. X=40.
Only one species known, NE Asia： NE China, Korea, N Japan, E Russia.
Plants of this genus look very close to the genera CORNOPTERIS Nakai and ATHYRIUM Roth morphologically, but there are some differences between them. Different from the former in lacking fleshy hornlike process at the base of costules adaxially; sori small, round or elliptic, never linear; not succulent plants, laminae papery when dried, greenish brown; Basal chromosome number X=40; occurs in the temperate zone, but not tropical and subtropical zone of Asia. Different from the typical ATHYRIUM plants in that rhizomes not erect, fronds moderately stout, standing on the long-creeping rhizomes in two rows; stipe not beak-shaped at base; laminae deltoid-ovate, basal pair of pinnae nearly as long as the second pair, all the lower pinnae petioled; bearing unicellular short hairs and thin, long, pale-brown multicellular hairs on the abaxial surface of raches, costae, especially costules; sori exindusiate. Molecular data shows that it is more close to the genus CORNOPTERIS.
Neoathyrium crenulatoserrulatum (Makino) Ching et Z. R. Wang in Acta Phytotax. Sin. 20(1):77, t. 1, 2. 1982.
新蹄盖蕨 xin ti gai jue
Athyrium crenulatoserrulatum Makino. ── Cornopteris crenulatoserrulata Nakai in Bot. Mag. Tokyo 45:95. 1931. as syn.; Phegopteris crenulatoserrulata Makino in Bot. Mag. Tokyo 17:78. 1903; Dryopteris crenulatoserrulata C. Chr.; Pseudoathyrium crenulatoserrulatum Nakai; Cornopteris coreana Nakai; Cornopteris crenulatoserrulata form. coreana H. Ito; Athyrium koryense C. Chr.; Dryopteris austro-ussuriensis Komarov; Phegopteris austro-ussuriensis Kom.; Athyrium austro-ussuriense Fomin.
Medium-sized terrestrial. Rhizomes moderately stout, creeping, sparsely clothed with brownish ovato-lanceolate, membranous big scales at apex and stipe bases; scales up to 1 cm or longer. Fronds distant; stipe nearly as long as or a little longer than lamina, 40--60 cm, the lower part moderately stout, not beak-shaped at base, up to 7--9 mm in diameter; laminae deltoid-ovate to ovato-oblong, 25--70 cm long, 20--60 cm broad, acuminate at apex, broadly cuneate or somewhat cordate at base, 2-pinnate, pinnules pinnatipartite; pinnae 10--15 pairs, broadly lanceolate or oblong-lanceolate, lower pinnae with 2--10 mm long short petioles, subopposite, ascendant, the basal 2 pairs of pinnae the largest, 10--32 cm long, 4--8 cm broad, acuminate at apex, slightly narrowed, broadly cuneate or near truncate at base, pinnate, pinnules pinnatipartite, upper pinnae gradually small, lanceolate, alternate, gradually to be sessile; pinnules 8--20 pairs, catadromic, lanceolate, 1--4 cm long, 5--15 mm broad, acuminate at apex, broadly cuneate at base, subsessile, the lowers subopposite, the uppers alternate, patent or slightly ascendant, pinnatipartite; ultimate segments 5--10 pairs, oblong, obtuse-rounded at apex, joining each other by narrow wing at base, dully toothed at margin. Veins unclear adaxially, visible abaxially, pinnate, midrib slightly tortuous, lateral veins forked. Laminae papery when dried, greenish-brown adaxially, green abaxially; lacking fleshy hornlike process at the base costules adaxially; raches, costae and costules on the abaxial surface bearing pallid unicellular short hairs and pale-brow multicellular long hairs. Sori round or elliptic, exindusiate, dorsal on the middle of veinlet. Spores bilateral, perispore surface with fold-like processes. 2n=80@.
Under subalpine mixed forests or grassland; 800--1000 m. Heilongjiang, Jilin, Liaoning, Shaanxi, Henan [Korea, N Japan, E Russia]. Type from Japan.
11. DRYOATHYRIUM Ching in Bull. Fan Mem. Inst. Biol. Bot. 11:39. 1941.
介蕨属 jie jue shu
PARATHYRIUM Holtt.; DEPARIA Sect. DRYOATHYRIUM (Ching) Kato; LUNATHYRIUM Sect. DRYOATHYRIUM (Ching) H. Ohba.
Terrestrial medium-sized plants. Rhizomes moderately stout, long-creeping, ascending or suberect, dictyostele. Fronds distant or approximate; stipes long, clothed with brown or brown, ovate to lanceolate scales at bases, inside 2 vascular bundles present, upward conjoint as U-shaped; laminae oblong or ovato-oblong, acuminate at apex, pinnate to 2-pinnate, ultimate pinnules pinnatipartite; pinnae or pinnules alternate, sessile, symmetrical, rotund-cuneate at base, pinnatifid to pinnatipartite at margin, segments oblong or rectangle, pinnules or segments joining with costae or costules by narrow wings; raches, costae and costules grooved adaxially, the grooves of raches or costae are not open to the grooves of costae or costules; usually bearing brown or dark brown, vermiculate glandular-hairs consisted of 1--3(--4) row of cells on costae, costules and midribs. Veins free, lateral veins simple or forked. Laminae herbaceous or papery when dried. Sori round, oblong, J-shaped or horseshoe-shaped, dorsal or lateral on the middle of veinlet; indusia same in shape with sori, membranous, entire, erose at margin, lacerate or ciliated, persistent. Spores bilateral, perispore surface with folds or spines, baculate ornamentation. X=40.
About 20 species. In the East Hemisphere, mainly in temperate and subtropical zone, some species can extend to tropical N Africa, Madagascar. East to Japan, Korea, E Russia, West to NW Himalayas. 14 species known in China, Mt. Qinling, W Hubei, Sichuan and NE Yunnan are the distribution center of the genus.
This genus is taxonomically close to ATHYRIUM, and more close to LUNATHYRIUM. But rhizomes in this genus long-creeping or ascending, fronds approximate, stipes not swollen as spindle and beak-shaped at base; bearing brown or dark brown, vermiculate glandular-hairs consisted of 1--3(--4) row of cells on the costae, costules or midribs, those are different from the genera ATHYRIUM and LUNATHYRIUM.
1a. Sori oblong or short-linear, in some cases curved; indusia oblong, lunate, J-shaped or horseshoe-shaped.
2a. Laminae 3-pinnate, pinnules petioled, pinnatilobate to pinnatipartite at margin, lateral veins pinnate; Jilin (Changbai Shan)
1. D. pterorachis
2b. Laminae pinnate-pinnatifid, pinnules entire or toothed at margin, lateral veins simple or forked.
3a. Laminae ovate, pinnae petioled; densely bearing multicellular hairs on the stipes and raches; spore surface with long spine-like processes. Hunan, Sichuan, Yunnan
2. D. erectum
3b. Laminae oblong, pinnae sessile; sparsely bearing vermiculate hairs consisted of 23 row of cells on stipes and raches; spore surface with fold processes.
4a. Segments entire (occasionally repand), lateral veins simple; sori mostly short-linear (in some cases slightly curved), occasionally J-shaped. Shaanxi and Sichuan
3. D. chinensis
4b. Segments toothed, lateral veins 2- to 3(4)-forked; sori horseshoe-shaped, J-shaped, oblong, elliptic etc.
5a. Laminae sparsely bearing brown spine-like thick hairs adaxially, lateral veins usually forked; Sichuan
4. D. setigerum
5b. Laminae subglabrous, bearing no spine-like thick hairs adaxially, lateral veins usually 2- to 4-forked.
6a. Rhizomes ascending or suberect; sori usually narrowly oblong, lunate or J-shaped; NE, N, NW and N Henan China; Korea and Japan
5. D. coreanum
6b. Rhizomes short-creeping; sori short-oblong, J-shaped or horseshoe-shaped; Shaanxi, Gansu, Fujian, Henan, Hubei, Hunan, Sichuan and Yunnan
6. D. henryi
1b. Sori mostly round or elliptic; indusia round-reniform, horseshoe-shaped, J-shaped or lunate.
7a. Laminae pinnate, pinnae pinnatipartite.
8a. Sori elliptic or rotund, indusia lunate, lateral or horseshoe-shaped; Shaanxi and Gansu
7. D. confusum
8b. Sori round or rotund, indusia round-reniform, occasionally horseshoe-shaped.
9a. Stipes and raches slightly bearing dark-brown ovato-lanceolate scales, segments crenated at margin, lateral veins forked to 3(4)-forked; Sichuan and Yunnan
8. D. stenopteron
9b. Stipes and raches clothed with quite a number of blackish brown, lustrous, broadly lanceolate scales, segments entire or toothed at margin; lateral veins forked, occasionally simple or 3-forked.
10a. Pinnae narrowly elliptic or elliptic, pinnae pinnatisect, segments uneven, the basiscopic one longer than the acroscopic one, often as falcate; Sichuan (Emei Shan)
9. D. falcatipinnulum
10b. Pinnae lanceolate, pinnatipartite, segments even, oblong; S Shaanxi, Taiwan, Sichuan and Yunnan
10. D. unifurcatum
7b. Laminae 2-pinnate-pinnatifid to -pinnatipartite.
11a. Pinnules more or less connate with costae at bases , sessile.
12a. Laminae thick herbaceous, pinnule subquadrangular at base, pinnatifid divided within 1/2, segments entire or repand-toothed; Shaanxi, Gansu, Jiangsu, Anhui, Zhejiang, Jiangxi, Fujian, Hubei, Hunan, Guangdong, Guangxi, Sichuan and Yunnan
11. D. okuboanum
12b. Laminae thin-herbaceous, pinnule broadly cuneate at base, pinnatipartite divided up to more than 2/3, segments toothed; Fujian and Sichuan
12. D. viridifrons
11b. Pinnule not connate with costae at base, shortly petiolulate.
13a. ultimate segments rounded at apex, toothed at margin; Hunan, Sichuan, Guizhou, Yunnan and Xizang
13. D. boryanum
13b. ultimate segments truncate or rounded-truncate at apex, entire at margin; Guangdong, Guangxi, Guizhou and Yunnan
14. D. edentulum
1. Dryoathyrium pterorachis (Christ) Ching in Bull. Fan Mem. Inst. Biol. Bot. 11:81. 1941.
翅轴介蕨 chi zhou jie jue
Athyrium pterorachis Christ in Bull. Bot. Herb. Boiss. 4:668. 1896 (Type, isotype KYO!); Cornopteris pterorachis Tard.-Blot; Lunathyrium pterorachis Kurata; Deparia pterorachis Kato.
Rhizomes long-creeping, ascending at apex. Fronds subcaespitose; fronds 1.2--1.5 m; stipes 30--70 cm long, 5--6 mm thick at base, sparsely bearing brownish, broadly lanceolate scales, upward pale brown-stramineous, glabrate; laminae oblong, 50--70 cm long or longer, 25--40 cm broad at middle, acuminate at apex, slightly narrowed at base, 2-pinnate, pinnules shallowly lobed to pinnatipartite; pinnae ca. 16 pairs, alternate, subsessile, slightly ascendant, the basal pair shorter, narrowed toward base, the second pair of pinnae linear-lanceolate, up to 20 cm long, 3--4 cm broad, acuminate at apex, symmetrical and truncate at bases, pinnate; pinnules 20--25 pairs, narrowly oblong or lanceolate, 1.5--2.5 cm long, 4--6 mm broad, obtuse-rounded at apex, broadened and connate with costae at base, the sinuses between pinnules often V-shaped, pinnatilobate to pinnatipartite at margin; segments oblong or subtriangular, entire; the third pairs and upper pinnae similar to the second pair but gradually shortened, pinnules shallowly lobed or toothed at margin. Veins pinnate on segments, lateral veins 2--4 pairs, veinlets simple. Laminae herbaceous when dried, dark green, raches, sparsely bearing brown lanceolate small scales and scale-like vermiculate hairs on costae and costules. Sori oblong or horseshoe-shaped, cross on acroscopic veinlets, 1(--2) sori per segment; indusia elongated or horseshoe-shaped, brownish, thick-membranous, entire, persistent. perispore surface with fold-like processes. 2n=80.
Damp areas under coniferous forests; 800--1000 m. Heilongjiang, Jilin [Korea, E Russia, Japan]. Type from Korea.
2. Dryoathyrium erectum (Z. R. Wang) W. M. Chu et Z. R. Wang in Fl Reip. Pop. Sin. 3(2):288. 1999.
直立介蕨 zhi li jie jue
Athyriopsis erecta Z. R. Wang in Acta Phytotax. Sin. 20(2):237, t.1, f.2, 4. 1982 (Type PE!); Deparia erecta Kato; Lunathyrium chingii Z. Y. Liu; Dryoathyrium chingii W. M. Chu apud P. S. Wang et X. Y. Wang; Dryoathyrium articulatipilosum ("articulati-pilosum") Ching et W. M. Chu apud Y. T. Hsieh.
Rhizomes creeping, ascending at apex. Fronds caespitose; fertile fronds up to 55--113 cm; stipes 30--40(--68) cm long, 2--3(--4.2) mm thick, not beak-shaped bases , sparsely bearing brown, ovate or broadly lanceolate scales, upward brownish stramineous, glabrate; laminae narrowly ovate, 25--40(--48) cm long, 14--24 cm broad at middle, acuminate at apex, rotund-cuneate at base, pinnate-pinnatipartite; pinnae 5--8(--12) pairs, basal 1--6 pairs subopposite, the basal pair of pinnae nearly with the same size to the second pair of pinnae, shortly petiolulate, petiolules up to 1.5 cm, upward the pinnae alternate, ascendant or subpatent, 2--5 cm distant, falcate-lanceolate, the middle and lower pinnae 10--14.5 cm long, 2.5--4 cm broad, acuminate at apex, nearly symmetrical, truncate at base, basal pair of pinnae prominently narrowed and a little asymmetrical at base, pinnatipartite at margin; segments 10--15(--17) pairs, the basals subopposite, upward alternate, slightly ascendant, close each other, oblong or rectangle, 1--2 cm long, 6--11 mm broad, obtuse-rounded or subtruncate at apex, shallowly toothed at margin. Veins pinnate in segments, lateral veins 6--9 pairs, simple or forked, occasionally 3- or 4-forked. Laminae herbaceous when dried, greenish brown, raches and costae上 sparsely bearing brown lanceolate small scales and scale-like hairs. Sori oblong, occasionally short-linear, J-shaped or horseshoe-shaped, 1--3 mm, dorsal on the middle of veinlets, 1--7 pairs per segment, uniserial along each side of midrib; indusia oblong, occasionally reniform or horseshoe-shaped, the basal acroscopic one often hooked, brownish, membranous, erose at margin, persistent. Perispore surface with long spine-like processes. 2n=80@.
* In valley under forests, wet areas; (200--)1200--2500 m. Hubei, Hunan, Sichuan, Chongqing, Guizhou, NE Yunnan. Type from Sichuan (Emei Shan).
This species is very easily to be recognized by the perispore surface with long spine-like processes, which is unique in the genus.
3. Dryoathyrium chinense Ching in Fl. Tsinling. 2:101,214--215. 1974. (Type PE!)
中华介蕨 zhong hua jie jue
Rhizomes creeping, ascending at apex. Fronds subcaespitose; fertile fronds up to 80 cm; stipes up to 30 cm long, ca. 2 mm thick at base, sparsely bearing brown lanceolate scales, upward stramineous, glabrate; laminae oblong, 40--50 cm long, 20--25 cm broad at middle, acuminate at apex, narrowed at base, pinnate-pinnatipartite; pinnae ca. 15 pairs, alternate, subsessile, slightly ascendant, lanceolate, the middle and lower pinnae 12--14 cm long, 2--2.5 cm broad, long-acuminate at apex, nearly symmetrical, truncate at base, pinnatipartite at margin; segments 17--20 pairs, oblong, 1--1.2 cm long, 6--7 mm broad, obtuse-rounded or subtruncate at apex, entire or sparsely repand at margin; the upper pinnae similar to the lowers in shape, but gradually shortened upward, pinnatifid to pinnatipartite, segments oblong or rectangle, entire. Veins pinnate in segments, lateral veins 6--8 pairs, usually simple, occasionally forked. Laminae thick herbaceous when dried, dark green, raches and costae上 sparsely bearing brown lanceolate small scales and scale-like vermiculate hairs. Sori short-linear, in some cases slightly curved, occasionally J-shaped or horseshoe-shaped, dorsal on the acroscopic veinlets, 5 or 6 pairs per segment, biserial along two sides of costule; indusia elongated, sometimes lunate, occasionally J-shaped or horseshoe-shaped, dark-brown, membranous, entire, persistent. Perispore surface with lophate folds.
* In valley under forests, damp areas; 1000--2100 m. S Shaanxi, Chongqing, Sichuan. Type from Chongqing (Chengkou).
4. Dryoathyrium setigerum Ching ex Y. T. Hsieh in Bull. Bot. Res. 5(3):153. 1985 (Type PE!).
刺毛介蕨 ci mao jie jue
Rhizomes creeping, ascending at apex. Fronds caespitose; fertile fronds 50--60 cm; stipes 20--25 cm long, ca. 2 mm thick at base, densely clothed with brown lanceolate scales, upward stramineous, glabrate; laminae oblong, 30--35 cm long, 16--20 cm broad at middle, acuminate at apex, narrowed at base, pinnate-pinnatipartite; pinnae 10--14 pairs, the basals subopposite, upward alternate, sessile, subpatent, lanceolate, the middles 10--12 cm long, 2--2.2 cm broad, acuminate at apex, symmetrical, subtruncate at base, pinnatipartite at margin; segments oblong, 8--10 mm long, 4--5 mm broad, obtuse-rounded or truncate at apex, entire or repand-crenate at margin. Veins pinnate in segments, lateral veins ca. 5 pairs, veinlets simple or forked. fronds herbaceous when dried, green, raches and costae bearing brownish broadly lanceolate small scales and scale-like vermiculate hairs. Sori oblong, J-shaped or horseshoe-shaped, medial or basal on veinlet, 4--6 pairs per segment, biserial along two sides of costule; indusia elongated, J-shaped or horseshoe-shaped, brown, membranous, entire, persistent. Perispore surface with lophate folds.
* In valley under forests, damp areas; 400--2100 m. Zhejiang, Hunan, Chongqing, Sichuan, Guizhou. Type from Chongqing (Chengkou).
5. Dryoathyrium coreanum (Christ) Tagawa in Journ. Jap. Bot. 22:162. 1948.
朝鲜介蕨 chao xian jie jue
Athyrium coreanum Christ in Bull. Herb. Bioss. ser. 2. 2:827. 1902. (Syntype, lectotype by Z. R. Wang: Fourie s.n., 28 Jun. 1901, Korea, P! Isotype KYO!); Lunathyrium coreanum (Christ) Ching; Deparia coreana Kato; Athyrium decursivum Yabe; Dryoathyrium henryi Fu, S. H., pro parte; Athyrium henryi auct., non Diels, 1899; Lunathyrium henryi sensu Kurata, pro parte; Deparia henryi sensu Kato, pro parte; Lunathyrium ningshanense Ching.
Rhizomes short-creeping, ascending or suberect, clothed with brownish, membranous, lanceolate scales at apices and stipe bases. fronds approximate; fertile fronds 60--80(--95) cm; stipes 30--40(--47) cm long, 3(--4) mm thick, swollen and dark-brown at base, upward stramineous, scales gradually sparse; laminae a little longer than stipes or almost equal, oblong-ovate, 18--25 cm broad at middle, pinnatifid-acuminate at apex, pinnate, pinnae pinnatipartite to pinnatisect; pinnae 12--15(--18) pairs, the lowers subopposite, slightly shortly petiolulate or subsessile, upswept, basal 1--2 pairs of pinnae a little shortened, prominently narrowed toward their base, 4--6 cm apart from each other, the middle pinnae lanceolate, 10--15 cm long, (1.5--)3 cm broad, long-acuminate at apex, subtruncate at base, (the acroscopic side nearly parallel to costae, the basiscopic side slightly cuneate), pinnatipartite or subpinnatisect; segments 15(--20) pairs, the basal pair of segments of the lowest 2--3 pairs of pinnae shorter, the middle segments oblong, (0.8--)1--1.2(--2) cm long, (4--)5 mm broad, obtuse or obtuse-rounded at apex, joining each other by narrow wing at base, serrate at margin, prominently spaced each other. Veins visible on surfaces, pinnate in segments, lateral veins 6--8 pairs, 2- to 3-forked. Laminae herbaceous when dried, pale-green, sparsely bearing brown multicellular long hairs along raches and costae on the abaxial surface, hairs more short and sparse on the adaxial surface. Sori mostly narrowly oblong or lunate, in some cases hooked, dorsal on the acroscopic veinlets of forked lateral veins; indusia same with sori in shape, brownish, sparsely ciliated at margin. Spores bilateral, surface with a few long fold-like processes. 2n=160,(240).
In valley under forests; 700--1000 m. Heilongjiang, Jilin, Liaoning, Hebei, Henan, Shaanxi, Gansu [Korea, Japan]. Type from Korea.
This species is very close to Dryoathyrium henryi (Bak.) Ching, so it was often been treated as a synonym of the later. However, for D. coreanum, rhizomes are ascending or suberect, sori usually are narrowly oblong, lunate or J-shaped; occurring in the north of Qinling Mountains; mostly tetraploid. But for D. henryi, rhizomes are short-creeping, sori are short-oblong, J-shaped or horseshoe-shaped; occurring in the south of Qinling Mountains; hexaploid.
Dryoathyrium mcdonellii (Bedd.) Z. R. Wang [in Fl. Reip. Pop. Sin. 3(2):291. 1999. Synonyms: Athyrium McDonellii Bedd.; Cornopteris mcdonellii Tard.; Parathyrium mcdonellii Morto; Lunathyrium Mcdonellii Ching; Deparia Mcdonellii Kato.] is very similar to Dryoathyrium coreanum morphologically . But it occurs in NW Himalayan (India, Pakistan and Kashmir). Its ploidy has not been known. What is the real relationship among these three species, that is still waiting for further investigations.
6. Dryoathyrium henryi (Bak.) Ching in Bull. Fan Mem. Inst. Biol. Bot. 11:81. 1941.
鄂西介蕨 e xi jie jue
Aspidium henryi Bak. in Ann. Bot. 5:306. 1891. (Type K!); Athyrium henryi Diels; Lunathyrium henryi Kurata; Deparia henryi Kato, pro parte.
Rhizomes short-creeping, ascending at apex. Fronds subcaespitose; fertile fronds 50--95 cm; stipes 20--35 cm long, 3--4 mm thick at base, sparsely bearing dark-brown lanceolate scales, upward stramineous, glabrate; laminae oblong, 30--60 cm long, 20--25 cm broad at middle, acuminate at apex, slightly narrowed at base, pinnate-pinnatipartite; pinnae 12--18 pairs, alternate, subsessile, slightly ascendant, broadly lanceolate, the ones below lamina middle 12--20 cm long, 3--4 cm broad, caudate-acuminate at apex, bases nearly symmetrical, truncate or rotund-cuneate, pinnatipartite at margin; segments falcate-oblong, 2--2.5 cm long, 6--8 mm broad, obtuse-rounded or shortly acute at apex, incised-toothed at margin; upper pinnae similar to the lowers, gradually shortened upward, pinnatifid to pinnatipartite, segments oblong or rhomboid, entire or serrate at margin. Veins在 segments 上为 pinnate, lateral veins 8--10 pairs, veinlets 2-3-forked. Laminae herbaceous when dried, dark green, raches and costae上 sparsely bearing brown broadly lanceolate small scales and scale-like vermiculate hairs. Sori short-oblong, in some cases curved or J-shaped, occasionally horseshoe-shaped, lateral on the acroscopic side of veinlets, rarely cross over veinlets, 5--7 pairs per segment, biserial along two sides of costule; indusia elongated, occasionally J- or horseshoe-shaped, brown, membranous, lacerate as ciliate-like at margin, persistent. Spore perispore surface wit many rugate folds. 2n=240.
* Damp areas under broad leaf forest or shrubs; 1000--2000 m. S Shaanxi, SE Gansu, Fujian, S Henan, Hubei, Hunan, Sichuan, Chongqing, Guizhou, Yunnan. Type from Hubei (Badong).
7. Dryoathyrium confusum Ching et Hsu in Fl. Tsinling. 2:101,213--214. 1974 (Type PE!).
陕甘介蕨 shan gan jie jue
Rhizomes creeping, ascending at apex. Fronds approximate; fertile fronds 50--65 cm; stipes 20--25 cm long, ca. 2 mm thick at base, sparsely bearing brown ovato-lanceolate scales, stramineous upward, glabrate; laminae narrowly ovate, 30--40 cm long, 23--28 cm broad at middle, acuminate and pinnatifid at apex, narrowed and rotund-cuneate at base, pinnate-pinnatipartite; pinnae ca. 16 pairs, alternate, subsessile, subpatent, lanceolate, the basal pair shortened, second pair 10--13 cm long, 2.4--3 cm broad, somewhat caudate-acuminate at apex, nearly symmetrical, rotund-cuneate at base, pinnatipartite at margin; segments ca. 15 pairs, the basals opposite, upward alternate, patent, oblong or rectangle, 1--1.5 cm long, 5--7 mm broad, obtuse-rounded at apex, subdentate at margin; the third pair of pinnae a little larger, morphology same with the second pair, from the fourth pair upward the pinnae gradually shortened. Veins pinnate in segments, lateral veins usually forked, only the basal acroscopic one often simple. Laminae herbaceous when dried, pale green, raches, costae and costules sparsely bearing brown broadly lanceolate small scales and brown scale-like vermiculate hairs. Sori sparse, rotund or elliptic, dorsal on the base of acroscopic veinlets, uniserial along each side of midrib; indusia lunate, occasionally horseshoe-shaped, pale-brown, membranous, entire, persistent. Perispore surface with baculate or spine-like ornamentation.
* Damp areas under broad leaf forests; 1300--1500 m. CS Shaanxi, SE Gansu. Type from Gansu (Kangxian).
8. Dryoathyrium stenopteron (Bak.) Ching in Y. L. Zhang et al., Sporae Pterid. Sin. 226, f.70a, 228, t.44, f.19,23. 1976.
川东介蕨 chuan dong jie jue
Polypodium stenopteron Bak. in Journ. Bot. 229. 1888, non Bak. 1886; Nephrodium stenopteron Diels; Dryopteris dielsii C. Chr.; Dryoathyrium dielsii Ching.
Rhizomes creeping, ascending at apex. Fronds approximate; fertile fronds 55--85 cm; stipes 20--40 cm long, 2--2.5 cm thick at base, sparsely bearing brown narrowly lanceolate scales, upward stramineous, glabrate; laminae ovato-oblong, 25--45 cm long, 18--30 cm broad at middle, acuminate and pinnatifid at apex, narrowed and rotund-cuneate at base, pinnate-pinnatipartite; pinnae 8--12 pairs, the basals subopposite, upward alternate, shortly stalked or sessile, sub-patent, broadly lanceolate, the middles 12--18 cm long, 2.5--4 cm broad, caudate at apex, asymmetrical at base, the acroscopic side narrowly cuneate or subtruncate, pinnatipartite at margin; segments 15--18 pairs, the basals nearly opposite, upward alternate, falcate-oblong, the basal pair shortened, the second and third pairs 1.2--2 cm long, 4--8 mm broad, obtuse-rounded or shortly acute at apex, crenulate at margin; pinnae from middle upward gradually shortened, pinnatifid to pinnatipartite, segments subentire or slightly crenate. Veins pinnate in segments, lateral veins 3- to 4-forked, occasionally forked. Laminae herbaceous when dried, green. Raches, costae and costules sparsely bearing brown lanceolate small scales and brown, scale-like vermiculate hairs. Sori small, round, dorsal on middle of veinlet, uniserial along each side of midrib; indusia small, round-reniform, brown, membranous, entire, persistent. Perispore surface with baculate and spine-like ornamentation.
* Damp areas under evergreen broad leaf forests or shrubs; 500--2200 m. Hubei, Hunan, Sichuan, Guizhou, Yunnan. Type from Hubei (Badong).
9. Dryoathyrium falcatipinnulum Z. R. Wang in Acta Phytotax. sin. 20:238--240, f.1. 1982 (Type PE!).
镰小羽介蕨 lian xiao yu jie jue
Rhizomes creeping. Fronds approximate; fertile fronds 50--60 cm; stipes 24 cm long, 2--3 mm thick at base, sparsely bearing blackish brown broadly lanceolate scales, upward stramineous, glabrate; laminae ovato-oblong, 30--34 cm long, 17--21 cm broad, acuminate at apex, rounded at base, pinnate, pinnae pinnatipartite, or nearly 2-pinnate; pinnae 12--15 pairs, the basal 1 or 2 pairs the largest, opposite, subsessile, the rests alternate, narrowly elliptic or elliptic, the lowers 10--14 cm long, 2.5--3.5 cm broad, long-acuminate at apex, narrowed, cuneate and asymmetrical at base, pinnatipartite to nearly pinnatisect; segments 14--16 pairs, the lowest 1 or 2 pairs prominently shortened, of the rest segments, the basiscopics longer than the acroscopics, the acroscopics oblong, 1.2--1.8 cm long, ca. 5 mm broad, rounded at apex, toothed at margin, the basiscopic one falcate-lanceolate, up to 3.2 cm long, ca. 6 mm broad, acuminate at apex, toothed at margin. Veins pinnate in segments, lateral veins 4--20 pairs, 2-3-forked, rarely simple. Laminae herbaceous when dried, dark green, raches, costae and costules sparsely bearing short multicellular brown hairs. Sori small, near round-reniform, dorsal on middle of veinlet, uniserial along each side of midrib; indusia small, reniform, dark-brown, membranous, entire, fugacious. Perispore surface with papillate or spine-like ornamentation.
* Under forests on mountain slopes; 700--1100 m, rare. Sichuan. Type from Sichuan (Emei Shan).
There are some plants in between this species and Dryoathyrium unifurcatum. Its hybrid origin can be estimated.
10. Dryoathyrium unifurcatum (Bak.) Ching in Bull. Fan Mem. Inst. Biol. Bot. 11:81. 1941.
峨嵋介蕨 e mei jie jue
Nephrodium unifurcatum Bak. in Journ. Bot. 26:228. 1888. (Type K!, Isotype KYO!); Dryopteris unifurcatum C. Chr.; Athyrium unifurcatum C. Chr.; Parathyrium unifurcatum Holtt.; Lunathyrium unifurcatum Kurata; Deparia unifurcata Kato; Aspidium pandiforme Christ; Dryopteris tosensis Kodoma; Dryoathyrium jinfoshanense Ching et Z. Y. Liu
Rhizomes long-creeping. Fronds distant; fertile fronds 45--95 cm; stipes 20--40 cm long, 2--3 mm thick at base, sparsely bearing blackish brown broadly lanceolate or linear scales, upward stramineous, glabrate; laminae ovato-oblong, 25--55 cm long, 20--28 cm broad at middle, acuminate and pinnatifid at apex, slightly narrowed at base, pinnate-pinnatifid; pinnae 12--14 pairs, the basals subopposite, upward alternate, subsessile, ascendant, lanceolate, the middles 13--16 cm long, 3--4 cm broad, acuminate at apex, narrowed and round-truncate at base, pinnatipartite at margin; segments 12--15 pairs, oblong, the basal pair shortened, the rests 1.5--2.5 cm long, 6--8 mm broad, obtuse-rounded or truncate at apex, entire at margin, pinnae from middle upward gradually shortened, pinnatipartite to pinnatipartite, segments oblong or near quadrangular, obtuse-rounded or truncate at apex. Veins pinnate in segments, lateral veins forked, occasionally 3-forked. Laminae herbaceous when dried, pale-green, raches, costae and costules上 sparsely bearing brown lanceolate small scales and brown, scale-like vermiculate hairs. Sori small, round, dorsal on veinlet middle, uniserial along each side of midrib; indusia small, round-reniform, red-brown, membranous, entire, persistent. Spores perispore surface with baculate or spine-like ornamentation. 2n=80@,160; "n"=2n=120(3X apo.).
Under forests in mountains, damp areas streamside; 200--2800 m. S Shaanxi, Zhejiang, Taiwan, Hubei, Hunan, Sichuan, Chongqing, Guizhou, Yunnan [Japan]. Type from Sichuan (Emei Shan).
11. Dryoathyrium okuboanum (Makino) Ching in Acta Phytotax. Sin. 10:303. 1965.
华中介蕨 hua zhong jie jue
Athyrium okuboanum Makino in Bot. Mag. Tokyo 13:16. 1899; Dryopteris okuboana Koidz.; Athyrium viridifrons Makino f. okuboanum Makino; Athyrium unifurcatum C. Chr. var. okuboanum H. Ito; Dryoathyrium viridifrons (Makino) Ching var. okuboanum Ching apud S H Fu; Lunathyrium unifurcatum Kurata var. okuboanum Kurata; Lunathyrium okuboanum Sugimoto; Deparia okuboana Kato.
Rhizomes creeping, ascending at apex. Fronds sub-caespitose; fertile fronds up to 1.2 m; stipes 30--50 cm long, 3--5 mm thick at base, sparsely bearing brown lanceolate scales, upward stramineous, glabrate; laminae broadly ovate or ovato-oblong, 30--80 cm long, 25--40 cm broad at middle, pinnatifid, acuminate at apex, rotund-cuneate and narrowed at base, 2-pinnate-pinnatilobate to 2-pinnate-pinnatipartite; pinnae 10--14 pairs, alternate, shortly petiolulate or nearly sessile, the basal pair a little shortened, oblong-lanceolate, 20--28 cm long, 5--9 cm broad, acuminate at apex, narrowed toward base, pinnate; pinnules 12--16 pairs, the basals subopposite, upward alternate, sessile, patent, the basal pair smaller, oblong, 1--1.2 cm long, ca. 5 mm broad, obtuse-rounded at apex, nearly symmetrical, broadly cuneate, decurrent and connate with narrow wings of costules at base,, shallowly lobed to pinnatipartite at margin, segments oblong, obtuse-rounded at apex, entire at margin. Veins pinnate in segments, lateral veins 2--4 pairs, simple. Laminae thick-papery when dried, grass green, raches, costae and costules上 sparsely bearing brownish broadly lanceolate small scales and scale-like vermiculate hairs. Sori round, dorsal on veinlets, usually 1 sorus per segment, occasionally 24 sori per segment; indusia round-reniform or somewhat horseshoe-shaped, greenish brown, membranous, entire, persistent. Perispore surface with baculate or spine-like ornamentation. "n"=2n=120(3X, apo).
In valley under forests, forest margin or wet areas streamside; Near sea level to 2500 m. S Shaanxi, SE Gansu, Jiangsu, Anhui, Zhejiang, Jiangxi, Fujian, Henan, Hubei, Hunan, Guangdong, Guangxi, Sichuan, Guizhou, Yunnan [Japan]. Type from Japan.
12. Dryoathyrium viridifrons (Makino) Ching in Bull. Fan Mem. Inst. Biol. Bot. 11:81. 1941.
绿叶介蕨 lu ye jie jue
Asplenium viridifrons Makino in Bot. Mag. Tokyo 6:46. 1892, nom. nud. (Type TI! Isotype MAK!); Athyrium viridifrons Makino in Bot. Mag. Tokyo 13:15.1899; Athyrium henryi Diels var. viridifrons Ohwi; Cornopteris viridifrons Tard.-Blot; Parathyrium viridifrons Holtt.; Lunathyrium viridifrons Kurata; Deparia viridifrons Kato.
Rhizomes creeping, moderately stout. Fronds approximate; fertile fronds up to 1.2 m; stipes 35--55 cm long, 3--4 mm thick at base, sparsely bearing brownish broadly lanceolate scales, upward stramineous, glabrate; laminae oblong, 40--65 cm, acuminate at apex, almost not narrowed and rotund-cuneate at base, 2-pinnate, pinnules pinnatipartite; pinnae 8--10 pairs, alternate or the basals subopposite, petioled, ascendant, the basal pair of pinnae oblong, 22--30 cm long, 10--15 cm broad at middle, acuminate at apex, truncate at base, pinnate; with pinnules 12--14 pairs, alternate, subsessile, nearly sessile, subpatent, lanceolate, the middle pinnae 6--9 cm long, 1.5--3.2 cm broad, acuminate at apex, somewhat cuneate at base, pinnatipartite at margin; segments 10--12 pairs, alternate, ascendant, rectangle, 6--12 mm long, 2.5--3.5 mm broad, obtuse-rounded at apex, dentate at margin. Veins pinnate in segments, lateral veins simple or forked. Laminae herbaceous when dried, green, raches, costae and costules sparsely bearing brownish lanceolate small scales and scale-like vermiculate hairs. Sori small, round or rotund, dorsal on veinlets, 1--3 pairs per segment, indusia round-reniform, brown, membranous, subentire, persistent. Perispore surface with many folds, as irregularly rugate. 2n=80.
Under forests or in forest margin; 300--2100 m. Zhejiang, Jiangxi, Fujian, Hunan, Sichuan, Guizhou, Yunnan [Korea, Japan]. Type from Japan.
Outline of this species is very similar to Dryoathyrium okuboanum (Makino) Ching, but laminae green when dried; pinnae broader; pinnules acuminate at apex, deeply pinnatipartite, cut almost to costae, segments serrate at margin, can be distinguished.
13. Dryoathyrium boryanum (Willd.) Ching in Bull. Fan Mem. Inst. Biol. Bot. 11:81. 1941.
介蕨 jie jue
Aspidium boryanum Willd. Sp. Pl. 4:285. 1810. (Type, isotype P!); Lastrea boryana Moore; Nephrodium boryanum Bak.; Dryopteris boryana C. Chr.; Athyrium boryanum Tagawa; Ctenitis boryana Cop.; Cornopteris boryanum Tard.-Blot; Parathyrium boryanum Holtt.; Lunathyrium boryanum H. Ohba; Deparia boryana Kato; Aspidium divisum Wall., nom. nud. (Syntype K!); Nephrodium divisum (Wall.) Hook.; Polypodium subtripinnatum Clarke; Dryopteris subfluvialis Hay.; Deparia subfluvialis Kato.
Rhizomes creeping, ascending at apex. Fronds subcaespitose; fertile fronds 1.2--2 m; stipes 40--95 cm long, up to 1 cm thick at base, sparsely bearing dark-brown subulate lanceolate scales, upward pale brown-stramineous, glabrate; laminae broadly ovate, 80--105 cm long, 60--85 cm broad at middle, acuminate at apex, narrowed and rotund-cuneate at base, 2-pinnate, pinnules pinnatipartite; pinnae 12--15 pairs, alternate, petioled, slightly ascendant, oblong-lanceolate, the basal pair up to 40 cm long, ca. 15 cm broad at middle, acuminate at apex, symmetrical, truncate at base, pinnate; pinnules 14--16 pairs, alternate, petioled, patent, broadly lanceolate, 6--10 cm long, 1.5--2 cm broad at middle, acuminate at apex, symmetrical, truncate at base, pinnatipartite at margin; segments ca. 12 pairs, suboblong, 7--10 mm long, 2.5--3.5 mm broad, obtuse-rounded at apex, crenate at margin. Veins pinnate in segments, lateral veins simple or forked. Laminae herbaceous when dried, yellow-green, sparsely bearing short pale hairs adaxially, raches, costae and costules sparsely bearing brown lanceolate small scales and scale-like vermiculate hairs. Sori small, round, dorsal on the middle of lateral vein or at the forfication of forked veinlets, 3--5 pairs per segment, biserial along two sides of costules; indusia round-reniform, brown, membranous, subentire, frequently abortive or fugacious. Perispore surface with stria-like folds. 2n=80.
Damp areas streamside under evergreen forests; 500--3300 m. Shaanxi, Zhejiang, Fujian, Taiwan, Hunan, Guangdong, Hainan, Guangxi, Sichuan, Guizhou, Yunnan, SE Xizang [Nepal, Sikkim, India, Sri Lanka, Vietnam, Myanmar, Thailand, Malaysia, Philippines, Indonesia; Africa]. Type from India.
Kato (1984) separated Deparia subfluvialis (Hay.) Kato from D. boryanum (Willd.) Kato by "rhizomes erect". However, the distribution area of D. subfluvialis is S China (from Taiwan to SW China) overlapping with D. boryana (Willd.) Kato. Moreover, the real erect rhizomes have never been found in any specimens of these species in the herbaria PE, at most, the rhizomes are creeping or ascending at apex. So, D. subfluvialis is not able to be distinguished from D. boryanum.
14. Dryoathyrium edentulum (Kunze.) Ching in Bull. Fan Mem. Inst. Biol. Bot.11:81. 1941.
无齿介蕨 wu chi jie jue
Aspidium edentulum Kunze. in Bot. Zeit. 474. 1846; Lastrea edentula Moore; Nephrodium edentulum Bak.; Lastrea edentula Moore; Nephrodium edentulum Bak.; Dryopteris edentula O. Ktze.
Rhizomes creeping, ascending at apex. Fronds subcaespitose; fertile fronds up to 1.2 m; stipes 35--55 cm long, ca. 6 mm thick at base, sparsely bearing brown linear-lanceolate scales, upward brownish stramineous, glabrate; laminae broadly ovate, 50--65 cm long, 30--50 cm broad at middle, acuminate at apex, narrowed and rotund-cuneate at base, 2-pinnate-pinnatifid to-pinnatipartite; pinnae 10--15 pairs, alternate, petioled, subpatent, oblong-lanceolate, the lowers 20--30 cm long, 9--11 cm broad at middle, acuminate and somewhat caudate at apex, symmetrical, subtruncate at base, pinnate; pinnules 15--18 pairs, the lowers subopposite, upward alternate, shortly petiolulate or subsessile, patent, oblong, 5--5.5 cm long, 1.5--1.7 cm broad at middle, obtuse at apex, broadly cuneate at base, pinnatifid to pinnatipartite at margin; segments oblong, 6--8 mm long, ca. 5 mm broad, truncate or rounded-truncate at apex, entire, upper pinnae same with the lowers in shape, but gradually shortened upward, bipinnatilobate to pinnatipartite; segments rectangle or subsquare, rounded-truncate or subrotund at apex, entire. Veins pinnate in segments, lateral veins 4 or5 pairs, veinlets simple or forked. Laminae herbaceous when dried, grass green, raches, costae and costules sparsely bearing brown lanceolate small scales and scale-like vermiculate hairs. Sori small, round, dorsal on lateral vein or veinlet middle, 2--4 pairs per segment; indusia small, round-reniform, red-brown, membranous, entire, persistent. Spore perispore surface with spine-like ornamentation.
Under forests or in valley, damp areas; 500--2400 m. Guangdong, Guangxi, Guizhou, Yunnan [Vietnam, India, Myanmar, Indonesia]. Type from Indonesia (Java).
12. LUNATHYRIUM Koidzumi in Acta Phytotax. Geobot. 1:30. 1932.
蛾眉蕨属 e mei jue shu
LUNATHYRIUM Sect. LUNATHYRIUM H. Ohba.; DEPARIA Sect. LUNATHYRIUM Kato.
Medium-sized plants under forests. Rhizomes erect or ascending, clothed with big scales at apices; scales brownish, red-brown or dark brown, ovato-lanceolate or narrowly lanceolate, membranous, entire. Fronds caespitose; stipes stramineous, chestnut-red or only chestnut-coloured on the abaxial side, usually shorter than lamina, beak-shaped at base, swollen as flat, concave-convex spindle above base, with protuberant pneumatophores along two lateral ridges of the spindle-like base, clothed with scales similar to that on rhizomes, more or less bearing multicellular, tortuous, hyaline hairs; laminae oblong-lanceolate or oblong-oblanceolate, occasionally narrowly ovate, pinnatifid-acuminate at apex, more or less narrowed toward base, occasionally only the basal 1--2 pairs of pinnae a little shortened or the basal pair the longest, pinnate-pinnatipartite; pinnae lanceolate, narrowly lanceolate, oblanceolate or long-elliptic, acuminate at apex, bases truncate, often abruptly broadened, usually laterally symmetrical, the basal acroscopic segments of lower pinnae sometimes prominently enlarged as auricled, sessile, alternate or the lowers subopposite, patent, basal several pairs of pinnae usually gradually shortened towards the base, in some cases the basal pair shortened as small auriculate lobes, pinnatipartite to the wings of costae, segments numerous, close, occasionally spaced, oblong or rectangle, occasionally obtuse-triangular, rounded or obtuse, occasionally acute or near truncate at apex, serrate, crenate or subentire at margin; raches and costae grooved adaxially, the groove of rachis is not open to the groove of costae; midrib visible, protuberant, not grooved adaxially, lateral veinlets 4--6(--9) pairs, usually simple, occasionally forked, upswept, the basal pair of lateral veins extend to the margin above sinus. Laminae herbaceous when dried, green or greenish brown, raches and costae in both or one surfaces and veins on surfaces more or less bearing hyaline multicellular hairs, bearing hyaline triangular tooth-like cells at segment margin and the sinus between segments, in some cases also with multicellular hairs. Sori short-linear to elliptic, 0.5--2 mm, straight, lateral on the middle acroscopic side of veins; indusia usually narrowly lunate or elliptic, short-linear when young, straight, in some cases, J-shaped, horseshoe-shaped or doubled on the top of laminae or pinnae, or on the suprabasal veinlets of segments, clam shell-shaped when matured, papery or thick-membranous, firm, persistent, yellow-brown, entire, subentire or slightly erose, occasionally shortly ciliated at margin, sori parallel each other like pectinate. Spores bilateral, bean-elliptic or elliptic, perispore surface with continual or discontinued folds, rugate, papillate, verrucose or baculate processes. X=40.
About 20 species and numerous varieties and hybrids, except Lunathyrium acrostichoides (Sw.) Ching occurred in E North America, most species occur in Asia, especially in China. From mountains of W China, towards the north to Mt. Qinling, NE China, and Japan, Korea, E Russia, towards the west to W Himalayas, towards the east to the mountains of central and east of China. Under mountainous broad and coniferous forests or in shrubs, damp areas or streamside.
This genus is very natural and quite identical morphologically. Different from the genus ATHYRIIUM Roth in that lamina usually pinnate, pinnae pinnatipartite, pinna symmetrical at base, raches and costae grooved adaxially, the groove of rachis is not open to the groove of costae, raches, costae and veins usually more or less bearing multicellular hyaline hairs, indusia lunate, rarely horseshoe-shaped or J-shaped, usually firm, clam shell-shaped; from view of bearing multicellular trichomes the genus is very close to ATHYRIOPSIS Ching and DRYOATHYRIUM Ching, but different from the laters in rhizomes erect, rarely ascending, stipes usually far shorter than laminae, laminae narrowly lanceolate or oblanceolate, numerous pairs of lower pinnae gradually shortened, the lowest several pairs of pinnae broadened at base, their basal acroscopic segments often auricled but not shortened.
Key to the species and varieties
1a. Rhizomes ascending or short-creeping; only 3 pairs of the lower pinnae a little shortened, the basal pair of pinnae usually more than 3 cm; multicellular short hairs present at the sinus between segments; perispore surface with a few continuous low folds or discontinuous ridgy folds. NE and N China, NE Asia and E North America.
2a. Hairs thin, short, sparse at the abaxial surfaces of raches and costae, E North America
L. acrostichoides
2b. Hairs thick, long, rather densely at the abaxial surfaces of raches and costae, NE Asia.
3a. Segments close each other, serrulate at margin; lateral veins simple; laminae and pinnae narrowly lanceolate or lanceolate. NE and N China, Korea, E Russia, N Japan
1. L. pycnosorum
3b. Segments distant each other, with long teeth at margin; lateral veins usually forked; laminae and pinnae broadly lanceolate. Heilongjiang
1b. L. pycnosorum var. longidens
1b. Rhizomes erect, more pairs of lower pinnae gradually shortened or only the basal 2--3 pairs a little shortened; multicellular short hairs absent or present at the sinus between segments; perispore surface with rugose, rugate, saccate, papillate, baculate processes, or ridgy folds. In the South area from N China, Central and South Japan.
4a. Multicellular hairs more or less present at the sinus between segments.
5a. Only a few pairs of lower pinnae a little shortened, the basal pair of pinnae never shortened as auriculate, usually more than 2 cm, stipes usually more than 10--15 cm.
6a. Indusia bearing short glandular-hairs on the back. W Sichuan and NW Yunnan
2. L. auriculatum
6b. Indusia with no or bearing short glandular-hairs only on the lower part of back. Shanxi, Shaanxi, Ningxia, S Gansu, Henan, Hubei, Sichuan
3. L. giraldii
5b. Numerous pairs of lower pinnae gradually shortened towards the base, the basal pair of pinnae no more then 1--2 cm, often auriculate; stipes usually within 10--15 cm.
7a. Multicellular long hairs densely present on surfaces of segments.
8a. Spore surface with a few discontinuous low folds or rugose folds; raches and costae densely bearing multicellular long hairs and scale hair; hairs absent at margin of segments. NW Yunnan
4. L. hirtiraches
8b. Spore surface with rugate, saccate folds, or papillate, verrucose processes; raches and costae only sparsely with multicellular hairs; segments with multicellular long hairs at margin.
9a. Short hairs sparsely present on the back of indusia only at the lower part. W Sichuan and NE Yunnan
5. L. liangshanense
9b. Multicellular long hairs densely present on the whole back of indusia. NW Yunnan
5b. L. liangshanense var. sericeum
7b. Multicellular hairs sparsely present on surfaces of segments.
10a. Hairs on the abaxial surface of aches and costae thick and long. S Gansu, W Hubei, Sichuan, Yunnan and SE Xizang
6. L. sichuanense
10b. Hairs on the abaxial surface of raches and costae thin, short.
11a. Pinnae deltoid-lanceolate, segments thinner, densely closed, only 2--3 mm broad, truncate at apex. NW Yunnan
6b. L. sichuanense var. gongshanense
11b. Pinnae oblong-lanceolate, segments broader, spaced, 4--6 mm broad, rounded at apex. W Hubei and S Chongqing
6c. L. sichuanense var. jinfoshanense
4b. Multicellular hair absent at the sinus between segments.
12a. Perispore surface with continuous low folds or discontinuous rugose folds.
13a. Numerous pairs of lower pinnae gradually shortened towards the base, the basal pair of pinnae no more than 2 cm, often auriculate.
14a. Fertile fronds only ca. 30 cm; segments thinner, less than 2 mm broad, subentire at margin, pinnae and segments both close each other; multicellular hairs thicker and longer on the abaxial surface of raches and costae. Xizang (Yadong) and Sikkim
7. L. sikkimense
14b. Fertile fronds more than 60 cm; segments more than 4 mm broad, serrulate at margin, pinnae and segments both separate each other; multicellular hairs thin and short on the abaxial surface of raches and costae. SE Xizang and NW Yunnan.
15a. Fertile fronds 60--70 cm, middle pinnae of lamina ca. 7 cm, segments 13--16 pairs. SE Xizang
8. L. medogense
15b. Fertile fronds 120--130 cm, middle pinnae of lamina 14--16 cm, segments 120--130 cm, middle pinnae of lamina 14--16 cm, segments up to 20--25 pairs. NW Yunnan
8b. L. medogense var. weimingii
13b. Only 2--3 pairs of lower pinnae a little shortened towards the base, the basal pair of pinnae usually more than 3 cm.
16a. Multicellular hairs very short on the upper surface of fertile pinnae, unclear; hairs thin, short and sparse on the abaxial surface of raches and costae; indusia subentire. N, NW China, NW Henan and N Sichuan.
17a. Laminae oblong or obovate, the lower pinnae a little narrowed toward their bases, only the basal pair of segments a little broadened; segments deltoid-oblong, long as two times as broad, toothed at margin; veinlets frequently forked. Beijing(Miyun)
9c. L. vegetius var. miyunense
17b. Laminae narrowly oblong or oblanceolate, lower pinnae not narrowed toward base; segments oblong, nearly as broad as long, serrulate or subentire at margin; veinlets simple.
18a. Laminae herbaceous; indusia brown, membranous, distally open when sori at maturity. N China, Shaanxi, S Gansu, Henan and N Sichuan
9. L. vegetius
18b. Laminae near papery; indusia pale-brown, papery, smooth, still cover sori as conchiform at maturity. N Sichuan
9b. L. vegetius var. turgidum
16b. Multicellular hairs visible on the upper surface of fertile pinnae; hairs thick and long on the abaxial surface of raches and costae; indusia slightly erose or shortly ciliated at margin. SE Xizang, occasionally W Sichuan.
19a. Middle pinnae with 10--13 pairs of segments, W Sichuan
10c. L. acutum var. liubaense
19b. Middle pinnae with more than ca. 14 pairs of segments, SE Xizang and Himalayas.
20a. Segments obtuse or rounded at apex, serrulate at margin, spaced each other, SE Xizang
10. L. acutum
20b. Segments near truncate at apex , toothed at margin, densely close each other. Xizang(Baga, Mainling)
10b. L. acutum var. bagaense
12b. Perispore surface with rugate, saccate folds or papillate, verrucose or baculate processes.
21a. Only a few pairs of lower pinnae a little shortened towards the base, the basal pair of pinnae usually more than 2 cm, never auriform, stipes usually more than 10--15 cm.
22a. Spore surface densely with baculate or spine-like processes. Sichuan(Emei Shan)
11. L. emeiense
22b. Spore surface sparsely bearing rugate, saccate folds or papillate, verrucose processes. Sichuan, S Gansu, W Hubei and NW Yunnan.
23a. Lower pinnae narrowly deltoid-ovate or ovato-lanceolate, prominently broadened towards the base, their basal segments reflexed. Sichuan
12. L. ludingense
23b. Lower pinnae lanceolate, not broadened towards the base, their basal segments explanate.
24a. Segments subentire or crenulate at margin. Gansu, Hubei, Sichuan, Guizhou, Yunnan and Xizang
13. L. wilsoni
24b. Segments prominently toothed at margin.
25a. Pinnae narrowly triangular, or segments obtuse-triangular; segments with long teeth at margin; lateral veins frequently forked, at least on the basal segments of middle pinnae. NW Yunnan.
26a. Pinnae narrowly triangular, gradually broadened towards bases; segments oblong or narrowly ovate, close each other, toothed at margin; lateral veins only on the basal segments of middle pinnae sometime forked. NW Yunnan
13b. L. wilsoni var. incisoserratum
26b. Pinnae lanceolate or oblanceolate, not gradually broadened towards bases; segments obtuse-triangular, widely distant, with long teeth like lobed at margin,; lateral veins mostly 2-forked, occasionally 3-forked. NW Yunnan
13c. L. wilsoni var. habaense
25b. Pinnae linear-lanceolate, segments oblong; segments serrulate at margin; lateral veins simple. W Sichuan, occasionally S Gansu.
27a. Plant more than 80--100 cm high, hairs thin and short on the abaxial surface of raches and costae. Gansu and Sichuan
13d. L. wilsoni var. maximum
27b. Plant below 80 cm high, multicellular hairs thick and long on the abaxial surface of raches and costae. Sichuan and Xizang
13e. L. wilsoni var. muliense
21b. Numerous pairs of lower pinnae gradually shortened towards the base, the basal pair of pinnae no more than 1--2 cm, often auriculate; stipes usually within 10--15 cm.
28a. Multicellular hairs thin, short and sparse at the abaxial surfaces of raches and costae.
29a. Segments truncate at apex. Hubei
14. L. vermiforme
29b. Segments obtuse-rounded or obtuse at apex. Zhejiang to NW Yunnan
15. L. shennongense
28b. Multicellular hairs thick, long and rather dense at the abaxial surfaces of raches and costae.
30a. Segments truncate at apex. NW Yunnan
16. L. truncatum
30b. Segments rounded or obtuse, occasionally rounded-truncate at apex. SW to E China, CS Japan.
31a. Fertile fronds less than 30 cm; pinnae less than 12 pairs, less than 5 cm; segments 5--7 pairs. Yunnan
17. L. brevipinum
31b. Fertile fronds more than 30 cm; pinnae up to 20 pairs, more than 5 cm; segments more than 12 pairs. SW to E China, Central and S Japan.
32a. Fertile fronds up to more than 1 m, laminae linear-oblanceolate, pinnae more than 25--30 pairs. Yunnan, occasionally W Sichuan.
33a. Segments bigger, 8--9 mm long, ca. 4 mm broad, prominently toothed at margin. Sichuan and Yunnan
18. L. dolosum
33b. Segments smaller, ca. 5 mm long, ca. 3 mm broad, subentire at margin. NW Yunnan
18b. L. dolosum var. chinense
32b. Fertile fronds 30--70 cm, laminae oblong or oblanceolate, pinnae 20--25 pairs. Mainly E China, occasionally central China, and Yunnan [Japan].
34a. Stipes usually brown-reddish, multicellular hairs of stipes and raches crispate. Anhui, Taiwan, CS Japan
19. L. orientale
34b. Stipes stramineous, multicellular hairs of stipes and raches straight. Anhui, Zhejiang, Jiangxi, Hubei, Sichuan
19b. L. orientale var. jiulungense
1. Lunathyrium pycnosorum (Christ) Koidz. Acta Phytotax. Geobot. 1:31. 1932.
东北蛾眉蕨 dong bei e mei jue
Athyrium pycnosorum Christ in Bull. Herb. Boiss. ser. 2. 2:827. 1902. (Syntype, Lectotype P! Type fragment BM!); Deparia pycnosora Kato; Athyrium acrostichoides Diels, pro parte; Lunathyrium acrostichoides (Sw.) Ching; Lunathyrium pycnosorum var. acutum J. J. Chien; Lunathyrium changbaiense Ching et J. J. Chien; Lunathyrium shandongense J. X. Li et F. Z. Li.
1a. var. pycnosorum
东北蛾眉蕨 dong bei e mei jue
Rhizomes stout, ascending, densely clothed with brownish, membranous, broadly ovate to ovato-lanceolate big scales at apices and stipe bases. Fronds caespitose; fertile fronds (30--)40--70(--87) cm; stipes 8--20(--31) cm, bases chestnut-black, upward gradually changed to stramineous, occasionally chestnut-reddish, sparsely bearing brown, membranous, lanceolate scales; laminae broadly lanceolate to oblong-lanceolate, (24--)30--40(--59) cm long, (5--)8--12(--18) cm broad, acuminate at apex, pinnate, pinnae pinnatipartite; pinnae 18--25 pairs, only a few lower pairs gradually shortened downward, usually the basal pair of pinnae more than 2.5 cm, rarely shorter than this, middle pinnae narrowly lanceolate, (2.5--)4--5(--9) cm long, 0.7--1.5 cm broad, acuminate at apex, subtruncate at base, the lowers subopposite, upward alternate, patent, 1--3 cm apart from each other, pinnatipartite almost up to costae; segments (7--)12--15(--19) pairs, close, 4--6 mm long, occasionally up to 1 cm long, 2--3 mm broad, oblong, rounded or obtuse and crenate at apex, entire or more or less crenate at margin. Veins visible on surfaces, pinnate in segments, lateral veins ca. 5 pairs per segment, veinlets simple. Laminae herbaceous when dried, green, slightly bearing multicellular short hairs along raches, costae, midribs and lateral veins. Sori long-lunate to linear, 3--5 pairs per segment, almost as long as veinlets, 1--1.5 mm, often densely close each other at maturity; indusia same with sori in shape, pale-brown, pectinate array on segment, persistent. Spores bilateral, perispore surface with a few low folds. 2n=80,160.
Damp areas under mixed forests; 200--1000 m. Heilongjiang, Jilin, Liaoning, Hebei, Shandong [Korea, N Japan, E Russia]. Type from Korea peninsula.
American Lunathyrium acrostichoides (Sw.) Ching possesses the characters of rhizome ascending, multicellular short hair present at the sinus between segments, spore surface with low folds. Which are very similar to those of L. pycnosorum. But the former with thin, short and sparse hairs on raches and costae abaxially, for the later hairs are quite thick and long. And for the later plant shorter, sori densely closer each other than the former. They mostly may be a couple of sibling species with disjunct distribution between E Asia--N America. So called "Lunathyrium acrostichoides" from NE to SW China in Chinese literatures before are incorrect.
1b. var. longidens Z. R. Wang in Fl. Reip. Pop. Sin. 3(2):509, 1999. (Type PE!).
长齿蛾眉蕨 chang chi e mei jue
It differs from the typical variety in far bigger pinnae and segments, pinnae 2--2.5 cm broad, segments ca. 1 cm long, 4--5 mm broad; segments prominently toothed at margin; shorter and sparser multicellular hairs at the sinus between segments; lateral veins frequently forked.
*Damp areas under mixed forests. Heilongjiang (Daguokui Shan, Shangzhi).
2. Lunathyrium auriculatum W. M. Chu et Z. R. Wang ex Z. R. Wang in Acta Phytotax. Sin. 3(1):85, t. 2, f. 1--4. 1994 (Holotype PYU! Isotype PE!).
大耳蛾眉蕨 da er e mei jue
2a. var. auriculatum
大耳蛾眉蕨 da er e mei jue
Rhizomes moderately stout, erect, clothed with dark-brown, membranous, lanceolate scales at apices and stipe bases. Fronds caespitose; fertile fronds 80--90 cm; stipes 10--25 cm long, 4--7 mm thick, stramineous, sparsely bearing brownish, hyaline, multicellular hairs, bases densely clothed with thick, dark-brown, linear-lanceolate scales, upward gradually sparse; laminae oblong-lanceolate or lanceolate, 20--63 cm long, 7--18 cm broad, pinnatifid-acuminate at apex, a little narrowed or almost not narrowed toward base, pinnate-pinnatipartite; pinnae 12--22 pairs, middle pinnae lanceolate, 7--13 cm long, 1.5--2.5 cm broad, acuminate at apex, bases slightly broadened, subtruncate, alternate, patent, pinnatipartite, only a few pairs of lower pinnae a little shortened or almost not shortened, abruptly prominently broadened at base, subopposite; segments ca. 18 pairs, oblong, separated each other by space, ca. 8 mm long, ca. 4 mm broad, rounded-obtuse or obtuse at apex, joining each other by narrow wing at base, the basal acroscopic segments of basal pair of pinnae prominently elongated, longer than other segments one time, become narrow-deltoid-ovate, acute to short-acuminate at apex, and pinnatipartite or pinnatilobate. Veins visible abaxially, pinnate in segments, lateral veins 5--7 pairs, simple, sometimes forked in lower segments of pinnae, upswept. Laminae herbaceous or near papery when dried, raches, costae and veins bearing rather densely thick, multicellular long hairs abaxially, short multicellular hairs present at the sinus between segments. Sori oblong or lunate, 47 sori per segment, 1--2 mm, close each other; indusia same with sori in shape, in some cases J-shaped at apices of lamina or pinnae, brownish, ciliated or prominently erose at margin, there are some glandular-hair-like attachments on the back. Spores bilateral, perispore surface with rugate, rugate-saccate or papillate processes.
* Under alpine shrubs, sparsely shaded wet places; 2900--3600 m. W Sichuan, NW Yunnan. Type from Yunnan (Yulong Shan, Lijiang).
This species is similar to Lunathyrium giraldii in shape, but the basal acroscopic segments of the basal pair or several lower pairs of pinnae prominently enlarged, and pinnatilobate to pinnatipartite; raches and costae rather densely with thick, long multicellular hairs abaxially; indusia prominently ciliated at margin, there are some glandular-hair-like attachments on the back.
2b. var. zhongdianense Z. R. Wang in Fl. Reip. Pop. Sin. 3(2):511, 1999. (Type PE!).
中甸蛾眉蕨 zhong dian e mei jue
It differs from the typical variety, L. auriculatum var. auriculatum, in the basal acroscopic segments of lower pinnae only a little enlarged, only with teeth or long teeth at margin, not pinnatipartite or pinnatilobate.
* Under coniferous forests on mountain slopes; ca. 2800 m. NW Yunnan (Zhongdian).
3. Lunathyrium giraldii (Christ) Ching in Act. Phytotax. Sin. 9:71. 1964.
陕西蛾眉蕨 shan xi e mei jue
Athyrium Giraldii Christ in Nuov. Giorn. Soc. Bot. Ital. new ser. 4:91. 1897.{Syntype, lectotype by Z. R. Wang: G. Giraldi No.55 Luglio 1894, Meta de Huan-tou-san, Shen-si selt., P! R. P. Licenln 2672 fragment, BM! excl. syntype: Padre B. Zou No.67 Estate del 1894, Shen-si, P! [=Matteuccia struthiopteris (L.) Todaro], G. Giraldi in 1896, Shen-si, P! & Estate 1896, Ki-fan, Shen-si, K! & syntype fragment, BM! [= Lunathyrium vegetius (Kitagawa) Ching]}; Athyrium subsimile Christ (Type P!); Athyrium sargentii C. Chr.; Lunathyrium acrostichoides auct. pro parte.
Rhizomes erect or ascending, clothed with brown, membranous, ovato-lanceolate scales at apices and stipe bases. Fronds caespitose; fertile fronds (30--)50--70(--90) cm; stipes (6--)10--15(--28) cm long, 1.5--4 mm thick, stramineous, occasionally chestnut-brownish abaxially, bases clothed with rather dense scales, upward sparse, nearly bearing no scales, bearing sparse, thin, short multicellular hairs; laminae oblong-lanceolate or ovato-lanceolate, (25--)35--60(--75) cm long, 10--15(--28) cm broad, acuminate at apex, slightly narrowed at base, pinnate; pinnae pinnatipartite, (15--)20--25 pairs, the middle pinnae linear-lanceolate, 5--8(--14) cm long, 1--2 cm broad, acuminate at apex, bases broader, truncate, alternate, ascendant, 2--2.5 cm apart from each other, only a few pairs of lower pinnae a little shortened, subopposite, the basal pair 2--4.5 cm, not auriculate; segments 15--22 pairs, oblong, (4--)6(--10) mm long, 3--4 mm broad, obtuse-rounded or obtuse at apex, joining each other by narrow wing at base, subpatent, crenulate or subentire at margin, separated each other by 12 mm space, pinnae the basal pair longer. Veins visible abaxially, pinnate in segments, lateral veins 4--6(--7) pairs, simple. Laminae herbaceous or near papery when dried, green or greenish brown, raches and costae on the abaxial surface sparsely bearing hyaline multicellular short hairs, multicellular hairs present too at the sinus between segments. Sori oblong to long-lunate, 1--1.5 mm, 2--6 pairs per segment; indusia same with sori in shape, pale-brown, erose or somewhat ciliated at margin, with no or bearing short glandular-hairs only on the lower part of back, persistent. Spores bilateral, perispore surface rugate-saccate, papillate or verrucose processes. 2n=80.
* In valley under forests; 900--2900 m. Shanxi, Shaanxi, Ningxia, S Gansu, Henan, Hubei, Sichuan. Type from Shaanxi (Taibai Shan).
4. Lunathyrium hirtiraches Ching ex Z. R. Wang in Act. Phytotax. Sin. 32(1):87, t.3, f.1--2. 1994 (Holotype PE!).
毛轴蛾眉蕨 mao zhou e mei jue
Rhizomes erect, densely scaly at apices and stipe bases; scales dark-brown, membranous, ovato-lanceolate, up to 1.5 cm. Fronds caespitose; fertile fronds ca. 80 cm; stipes ca. 15 cm long, 3--4 mm thick, stramineous pale-brownish, densely scaly at bases, upward densely clothed with narrowly lanceolate scales and brownish multicellular long hairs; laminae narrowly oblong, ca. 65 cm long, ca. 20 cm broad, acuminate at apex, gradually narrowed toward base, pinnate-pinnatipartite; pinnae 20--30 pairs, the middle pinnae linear-lanceolate, up to 10 cm long, ca. 1.8 cm broad, alternate, ascendant, 2--3 cm apart from each other, acuminate at apex, bases near truncate, pinnatipartite, 5--6 pairs of the lowers gradually shortened towards the base, lanceolate, the basal pair auriculate, 1--2 cm, basiscopic side broader; segments 20 pairs or more, oblong, ca. 8 mm, rounded at apex, joining with broad wings of costae at base, toothed at margin, patent, close each other. Veins on the abaxial surface clearer, pinnate in segments, lateral veins 5--7 pairs, simple. Laminae herbaceous when dried, raches and costae bearing rather dense multicellular long hairs on surfaces, veins sparsely bearing brownish, hyaline multicellular long hairs too on surfaces, bearing a few short multicellular hairs at the sinus between segments. Sori oblong, medial on veinlets, 3--4 pairs per segment, ca. 1 mm; indusia same with sori in shape, brown, slightly bulgy, ciliate-lacerate at margin, persistent. Spores bilateral, perispore surface with a few low continuous folds.
Evergreen forest margin, streamside, damp area; 2400--3900 m. NW Yunnan [NW Myanmar]. Type from Yunnan (Gongshan to Fugong, Watershed of west bank of Nujiang River-Dulongjiang River).
This species similar to Lunathyrium dolosum in shape, but spore surface with a few continuous folds, with no auriculate lobe-like processes, densely bearing multicellular long hairs and scale-like hairs on stipes and raches, lamina bearing multicellular long hairs on surfaces, so can easily be distinguished.
5. Lunathyrium liangshanense Ching ex Z. R. Wang in Acta Phytotax. Sin. 32(1):89, t.4, f.1--4. 1994 (Holotype PE!).
凉山蛾眉蕨 liang shan e mei jue
5a. var. liangshanense
凉山蛾眉蕨 liang shan e mei jue
Rhizomes stout, ascending. Fronds caespitose; fertile fronds (50--)100 cm; stipes far shorter than lamina, ca. 15 cm long, 3--3.5(--5) mm thick, stramineous, densely clothed with brown, lanceolate scales and multicellular, tortuous, hyaline hairs at bases, upward sparsely; laminae oblong-lanceolate or oblanceolate, (40--)80(--90) cm long, (11--)16(--18) cm broad at middle, pinnatifid-acuminate at apex, bases narrowly cuneate, pinnate, pinnae pinnatipartite; pinnae(20--)25--30(--35) pairs, the middle pinnae lanceolate, (6--)10(--12) cm long, 1--2 cm broad, acuminate at apex, a little broadened at base, broadly cuneate or subtruncate, alternate, ascendant or subpatent, 2--3 cm apart from each other, numerous pairs of lower pinnae gradually shortened, 3--4 pairs of the basal pinnae less than 1 cm as deltoid auricles; segments (14--)20 pairs, oblong, (4--)8(--10) mm long, (2.5--)4(--5) mm broad, rounded at apex, joining with broad wings of costae at base, subentire or slightly crenate, close each other. Veins visible abaxially, pinnate in segments, lateral veins (4--)6--7 pairs, simple. Laminae herbaceous when dried, rather densely bearing tortuous, hyaline multicellular hairs on surfaces and at the segment margins and the sinus between segments. Sori short-linear, 3--6 pairs per segment, 1--2 mm, close each other; indusia same with sori in shape, curved or double on the apices of lamina and pinnae, brownish, pilose or almost glabrous, erose like or sparsely short-ciliated at margin. Spores bilateral, perispore surface with rugate, rugate-saccate or papillate processes.
* Streamside, under forests, damp area; 1900--2200 m. SW Sichuan, W Guizhou, Yunnan. Type from Sichuan (Liang Shan, Leibo).
This species is similar to Lunathyrium giraldii in shape, but numerous pairs of lower pinnae gradually shortened towards the base, basal pair of pinnae often auriculate, stipes shorter, densely bearing multicellular hairs all over, segments also with multicellular long hairs at margin, so it is easy to be recognized.
5b. var. sericeum Ching et Z. R. Wang ex Z. R. Wang in Acta Phytotax. Sin. 32(1):89, t.4, f. 5--6. 1994 (Holotype PE!).
绢毛蛾眉蕨 juan mao e mei jue
It differs from the typical variety in plant smaller, fertile fronds ca. 45 cm, middle pinnae 4--5 cm, densely sericeous on leaf surfaces, and with multicellular long hairs on indusia.
* Alpine mountains, NW Yunnan.
6. Lunathyrium sichuanense Z. R. Wang in Acta Phytotax. Sin. 32(1):87, t. 3, f.3--5. 1994 (Holotype PE!).
四川蛾眉蕨 si chuan e mei jue
6a. var. sichuanense
四川蛾眉蕨 si chuan e mei jue
Rhizomes erect, clothed with brownish, membranous, ovato-lanceolate scales at apices and stipe bases. Fronds caespitose; fertile fronds (30--)55--65(--105) cm; stipes far short than lamina, (4--)8--12(--24) cm long, (1--) 2--3(--4) mm thick, stramineous or slightly chestnut-coloured abaxially, bases densely scaly, upward sparse or nearly bearing no scales; laminae elliptic to oblanceolate, (24--)45--55(--80) cm long, (7.5--)14--18(--27) cm broad, pinnatifid-acuminate at apex, gradually narrowed toward base, pinnate-pinnatipartite; pinnae (14--)20--25(--27) pairs, the middle pinnae linear-lanceolate, (4--)8--10(--14) cm long, 1--1.8 cm broad, alternate, ascendant or patent, acuminate at apex, bases near truncate, (1--)2--4(--6) cm apart from each other, numerous pairs of lower pinnae gradually shortened, lanceolate, subopposite, the basal pair often shortened as auriform, 0.5--2 cm; segments (8--)12--18(--22) pairs, oblong, 4--7 mm long, ca. 4 mm broad, obtuse-rounded or obtuse at apex, joining with the narrow wings of costae at base, ascendant. Veins visible on surfaces, pinnate in segments, lateral veins 4--6 pairs, simple. Laminae herbaceous when dried, greenish brown, raches and costae bearing rather densely multicellular hairs abaxially, with multicellular short hairs at the sinus between segments. Sori oblong or short-linear, medial on veinlets, 1--2 mm; indusia same with sori in shape, glabrous or short hairs only present on the lower part above, ciliated-like at margin, persistent. Spores bilateral, perispore surface with rugate-saccate, rugate or papillate processes.
* Mountain slopes, or damp area streamside; 1400--3400 m. S Gansu, Sichuan, Chongqing, Guizhou, Yunnan, SE Xizang. Type from Sichuan (Hongxi).
This species is similar to Lunathyrium liangshanense in shape, but laminae sparsely hairy on surfaces, segments without multicellular long hairs at margin, at most occasionally with 1-2 short hairs between teeth; This species is also similar to L. shennongense, but bearing multicellular hairs at the sinus between segments, raches and costae densely with longer multicellular hairs abaxially. So it is easy to be recognized.
6b. var. gongshanense Z. R. Wang in Acta Phytotax. Sin. 32(1):88--89. 1994 (Holotype PE! Isotype KUN!).
贡山蛾眉蕨 gong shan e mei jue
It differs from the typical variety in raches and costae only with thin, short and sparse multicellular hairs abaxially, subglabrous, pinnae deltoid-lanceolate, segments thinner and densely close each other, 2--3 mm broad, truncate at apex.
* Under coniferous forests on mountain slopes; ca. 3500 m. Yunnan (Gongshan).
6c. var. jinfoshanense Z. R. Wang in Fl. Reip. Po. Sin. 3(2):511, 1999. (Type PE!)
金佛山蛾眉蕨 jin fo shan e mei jue
It differs from the typical variety in raches and costae only with thin, short and sparse hairs abaxially, subglabrous; segments rounded at apex, undulate-crenulate at margin.
* Under forests on mountain slopes; 1500--2100 m. Hubei, Chongqing. Type from Chongqing (Jinfo Shan, Nanchuan).
7. Lunathyrium sikkimense Ching in Acta Phytotax. Sin. 9:72. 1964 (Type, isotype PE!).
锡金蛾眉蕨 xi jin e mei jue
Rhizomes ascending. Fronds caespitose; fertile fronds only ca. 5 cm; stipes ca. 10 cm long, ca. 1.5 mm thick, densely scaly at base, upward glabrate, stramineous; laminae narrowly oblong-lanceolate, 30 cm long,7 cm broad, acuminate at apex, pinnate, pinnae pinnatipartite; pinnae ca. 16 pairs, 3--4 pairs of the lowers gradually shortened towards the base, the basal pair ca. 1.4 cm, subopposite, spaced, middle pinnae linear, 4 cm long, 8 mm broad, short-acuminate at apex, bases truncate, alternate, 1.2 cm apart from each other, upswept, pinnatipartite; segments 12--14 pairs, thin, close each other, rectangle or oblong, 3--4 mm long, less than 2 mm broad, obtuse or subtruncate at apex, entire. Veins visible abaxially, veinlets 4--5 pairs per segment, simple. Laminae papery when dried, brown, raches stramineous, raches and costae bearing more brown-red multicellular hairs abaxially. Sori small, broadly ovate or oblong, 2--4 pairs per segment; indusia same with sori in shape, entire, dark-brown. Spores bilateral, perispore surface rugose.
Under forests; 3000--4300 m. Xizang (Yadong) [Sikkim]. Type from Sikkim (Darjeeling).
It is the smallest species in the genus, its pinnae and segments are very smaller and thinner. It is easy to be recognized.
8. Lunathyrium medogense Ching et S. K. Wu in C. Y. Wu, Fl. Xizang. 1:120, t. 29, f. 1--3, pl. 6, f. 3--4. 1983 (Type PE!).
墨脱蛾眉蕨 mo tuo e mei jue
8a. var. medogense
墨脱蛾眉蕨 mo tuo e mei jue
Rhizomes moderately stout, erect, bearing remainder stipe bases, up to 5--6 cm thick, scaly at apices and stipe bases; scales big, red-brown or brown, membranous, ovato-lanceolate, up to 15 mm long, 3--4 mm broad. Fronds caespitose; fertile fronds 70--115 cm; stipes 10--30 cm long, 3--5 mm thick, stramineous, in some cases slightly chestnut-coloured, bases densely scaly, upward scales gradually sparse; laminae oblanceolate or oblong- oblanceolate, 48--85 cm long, 12--26 cm broad at middle, pinnatifid and short-acuminate at apex, gradually narrowed toward base, pinnate-pinnatipartite; pinnae 20--25 pairs, the lowers 5--7 pairs gradually shortened towards the base, subopposite, upwards alternate, patent or slightly ascendant, the basal pair of pinnae auriform, ca. 1 cm, ca. 4 cm apart from each other, middle pinnae narrowly lanceolate, 7--13.5 cm, acuminate or long-acuminate at apex, subtruncate or broadly cuneate, broadest at bases, 1.2--2 cm, up to 2.5 cm broad at bases, pinnatipartite almost to costae; segments of middle pinnae more than 15--20 pairs, patent, oblong, obtuse-rounded or obtuse at apex, broader, joining each other by narrow wing at base, separated each other by narrow space, prominently toothed at margin. Veins slightly visible on surfaces, pinnate in segments, lateral veins5--7 pairs, simple, upswept. Laminae herbaceous when dried, raches, costae and veins sparsely bearing thin, short multicellular hairs abaxially, sparsely bearing red-brown short thick hairs adaxially. Sori oblong or short-linear, occasionally J-shaped or horseshoe-shaped on the basal acroscopic side of the top segments of laminae or pinnae, 1--2 mm, 3--6 pairs per segment, close each other; indusia same with sori in shape, brownish, erose at margin or subciliated, persistent. Spores bilateral, perispore surface with a few low folds.
* Damp areas under broad leaf forests; 2800--4000 m. NW Yunnan, SE Xizang. Type from Xizang (Medog).
This species is very similar to the plants so-called Lunathyrium allantodioides (Bedd.) Ching [in Acta Phytotax. Sin. 9:72. 1964; Athyrium allantodioides Bedd., Ferns Brit. Ind. t. 221. 1867. (Type K!); Deparia allantodioides Kato] from N India, Sikkim, Bhutan, Nepal, Pakistan and Taiwan. The chromosome number reports from India were diploid or tetraploid, 2n=80 or 160. But the spores of the type specimens in K are unequal-sized, rounded, black abortive, which shows it should be a hybrid, Lunathyrium Χ allantodioides (Bedd.) Z.R. Wang (in Fl. Reip. Pop. Sin. 3(2):308, 1999.). The relationship between them is still waiting for further study.
8b. var. weimingii Z. R. Wang in Acta Phytotax. Sin. 32(1):89. 1994 (Holotype PE! Isotype PYU!).
维明蛾眉蕨 wei ming e mei jue
It differs from the typical variety in rhizome ascending, plant far bigger, up to 120--130 cm high, middle pinnae up to 14--16 cm, their segments up to 20--25 pairs.
* Under mixed or coniferous forests; 2500--3300 m. Yunnan. Type from Yunnan (Gongshan).
9. Lunathyrium vegetius (Kitagawa) Ching in Acta Phytotax. Sin. 9:71. 1964.
河北蛾眉蕨 he bei e mei jue
Athyrium pycnosorum Christ var. vegetius Kitagawa in Rep. First Sci. Exped. Manch. 4(2):72. 1935 (Isotype: PE!); Lunathyrium pycnosorum Koidz. var. vegetius Kurata; Athyrium acrostichoides Diels, pro parte; Lunathyrium acrostichoides Ching, pro parte.
9a. var. vegetius
河北蛾眉蕨 he bei e mei jue
Rhizomes erect, in some cases branched as clustered, densely clothed with red-brown or brown, membranous, broadly lanceolate big scales, long scale-hair and multicellular hairs at rhizome apices and stipe bases. Fronds caespitose; fertile fronds (40--)60--80(--150) cm; stipes (10--)20--25(--50) cm long, 1--3(--5) mm thick, stramineous, red-brownish or brown, grooved adaxially, subglabrous abaxially; laminae narrowly oblong or oblanceolate, (30--)35--60(--100) cm long, (12--)16--24(--26) cm broad at middle, acute at apex, pinnate-pinnatipartite; pinnae (15--)20 pairs, ca. 3 pairs of lowest pinnae a little shortened towards the base, (1.5--)2--3(--9) cm apart from each other, the basal pair 3--4 cm, subopposite, the middle pinnae lanceolate, (6--)9--13 cm long, 1--2 cm broad, acuminate at apex, bases broadly cuneate, alternate, ascendant, pinnatipartite; segments (10--)15--18(--20) pairs, oblong, ovato-oblong or obtuse-triangular, 5--7(--9) mm long, 3--7 mm broad at base, rounded-obtuse or obtuse-acute at apex, slightly ascendant, subentire or undulate-crenate at margin. Veins depressed adaxially, slightly convex abaxially, pinnate in segments, lateral veins 4--5 pairs, veinlets simple. Laminae papery when dried, greenish brown, sterile lamina bearing short multicellular hairs conspicuously adaxially, fertile fronds surfaces subglabrous, raches and costae with the same colour to stipes, sparsely bearing short hairs or subglabrous abaxially. Sori oblong, 2--4 pairs per segment; indusia lunate, ca. 2 mm, subentire, pale-brown, persistent. Spores bilateral, perispore surface with a few low folds. 2n=160@.
* In valley under forests, wet areas or streamside; 400--2600 m. Hebei, Shanxi, Shaanxi, Gansu, Shandong, Henan, NW Sichuan. Type from Hebei (Changshanyu).
9b. var. turgidum Ching et Z. R. Wang in Acta Phytotax. Sin. 32(1):91. 1994 (Holotype PE!).
壳盖蛾眉蕨 ke gai e mei jue
It differs from the typical variety in indusia pale-brown, papery, surface more smooth, still cover sori as conchiform at maturity, perispore surface rugose.
* Damp areas streamside under forests, 1400--2800 m. Chongqing, Sichuan. Type from Sichuan (Miyaluo, Li Xian).
9c. var. miyunense Ching et Z. R. Wang ex Z. R. Wang in Fl. Reip. Pop. Sin. 3(2):509, 1999. (Type PE!).
密云蛾眉蕨 mi yun e mei jue
It differs from the typical variety in lamina oblong or obovate; lower pinnae a little shortened toward base, the basal pair of pinnae prominently shortened; segments broader and longer, deltoid-ovate or oblong, obtuse at apex, prominently toothed at margin, widely separated each other; veins frequently forked. 2n=ca.160.
* Damp areas streamside under broad leaf forests; 1100--1200 m. Beijing (Wuling Shan, Miyun).
10. Lunathyrium acutum Ching in Acta Phytotax. Sin. 9:73. 1964 (Type, isotype PE!).
尖片蛾眉蕨 jian pian e mei jue
Lunathyrium tibeticum Ching; Lunathyrium bomiense Ching et S. K. Wu; Lunathyrium latibasis Ching; Deparia shikkimensis Nakaike et S. Malik.
10a. var. acutum
尖片蛾眉蕨 jian pian e mei jue
Rhizomes erect, densely scaly at apices and stipe bases; scales red-brown, membranous, oblong, acuminate at apex. Fronds caespitose; fertile fronds 30--80 cm; stipes 15--21 cm long, 2--2.5 mm thick, pale stramineous, densely scaly at bases, scales brown, lanceolate, ca. 1 cm, upward naked; laminae long-elliptic or oblong-lanceolate, 35--45 cm long, 10--15 cm broad at middle, acuminate at apex, narrowed toward base, pinnate-pinnatipartite; pinnae 15--20 pairs, slightly ascendant, sessile, separated by narrower spaces, the lowers subopposite, upwards alternate, only a few pairs of lower pinnae a little shortened, the basal pair of pinnae 2.5--4 cm, the second pair up to 6 cm, middle pinnae lanceolate, somewhat falcate, 7--9 cm long, 1.5--2 cm broad at base, 2--3 cm distant, acuminate at apex, symmetrical, truncate, broader at base, pinnatipartite cut to the narrow costal wings; segments 13--17 pairs, subpatent, spaced, oblong, obtuse or obtuse-rounded at apex, serrulate or subentire at margin, the basal pair bigger, ca. 1 cm long, ca. 4 mm broad, the superiors 7--8 mm long. Veins visible abaxially, lateral veins 5--7 pairs per segment, up to 9 pairs in the basal segments, veinlets simple, upswept. Laminae herbaceous when dried, pale greenish-brown, raches and costae bearing brown multicellular hairs abaxially, costules of segments bearing prominently sparse short thick hairs on surfaces, bearing no multicellular hairs at the sinus between segments. Sori lunate or short-linear, 3--5 pairs per segment, up to 6 pairs in the basal segments, upswept; indusia same with sori in shape, firm, brown, entire, persistent. Spores bilateral, perispore surface with rugose folds.
Under coniferous forests or in valley; 2700--4000 m. SE Xizang [N India, N Pakistan; W Himalayas]. Type from India.
10b. var. bagaense (Ching et S. K. Wu) Z. R. Wang in Fl. Reip. Pop. Sin. 3(2):305, 1999. (Type PE!).
巴嘎蛾眉蕨 ba ga e mei jue
Lunathyrium bagaense Ching et S. K. Wu. in C. Y. Wu, Fl. Xizang. 1:118, t. 29, f. 4--6. 1983. (Type PE!).
It differs from the typical variety in segments oblong, densely close each other, obtuse at apex, near truncate, dentate at margin.
* Under forests, ca. 3400 m. Xizang. Type from Xizang (Paka, Mainling).
10c. var. liubaense (Z. R. Wang) Z. R. Wang in Fl. Reip. Pop. Sin. 3(2):305, 1999. (Type PE!).
六巴蛾眉蕨 li ba e mei jue
Lunathyrium vegetius (Kitagawa) Ching var. liubaense Z. R. Wang in Acta Phytotax. Sin. 32:91. 1994 (Holotype PYU! Isotype CDBI!).
It differs from the typical variety in segment number of pinnae far fewer, only 10--13 pairs, so that it is easily recognized.
* About 3300 m. Sichuan (Liuba, Kangding).
The spore surface of this variety presents fold processes, morphology of this variety is in between L. acutum and L. vegetius. It is similar to L. acutum by multicellular hairs longer on the abaxial surface of raches and costae, multicellular hairs prominent too on the adaxial surface of fertile pinnae, indusia shortly ciliated at margin; however, segment number of its pinnae is far fewer, that is similar to L. vegetius. This variety occurs also in the locality between L. acutum and L. vegetius. Their relationship is waiting for further studying.
11. Lunathyrium emeiense Z. R. Wang in Fl. Reip. Pop. Sin. 3(2):510, 1999. (Type PE!).
棒孢蛾眉蕨 bang bao e mei jue
Fronds caespitose; fertile fronds 90 cm; stipes ca. 20 cm long, ca. 4 mm thick, nearly bearing no scales and hairs; laminae narrowly oblong-lanceolate, ca. 58 cm long, ca. 18 cm broad, acuminate at apex, gradually narrowed towards the base, pinnate-pinnatipartite; pinnae ca. 20 pairs, middle pinnae linear-lanceolate, ca. 10.5 cm long, 1.8--2 cm broad, long-acuminate at apex, broader and subtruncate at base, nearly lateral symmetrical, alternate, ascendant, pinnatipartite, 5--6 pairs of the lowers gradually shortened towards the base, the basal pair lanceolate, ca. 2 cm long, ca. 1 cm broad, patent, the acroscopic side far broader at base, ca. 5 cm apart from the second pair, upward pinnae gradually closer; segments ca. 15 pairs, oblong, obtuse-rounded at apex, joining each other by narrow wing at base, serrulate at apices and top margin, ascendant, separated each other by narrow space. Veins visible on surfaces, pinnate in segments, lateral veins 6--7 pairs, simple, occasionally forked, upswept. Laminae herbaceous when dried, greenish brown, raches, costae and veins on the abaxial surface occasionally with tiny multicellular hairs or subglabrous. Sori oblong to short-linear, 1--2 mm long, medial on veinlets, 2--4 pairs per segment; indusia same with sori in shape, brown, slightly erose at margin. Spores bilateral, perispore surface densely with baculate or thin long rugate processes.
* In shrub groves on mountain slopes, damp areas, 2000--2100 m. Sichuan (Emei Shan).
This species is very similar to Lunathyrium wilsoni, however the segments prominently toothed at margin, spore surface densely with baculate or thin long rugate processes.
12. Lunathyrium ludingense Z. R. Wang et L. B. Zhang in Acta Bot. Yunnan. 17(3):303--304, t.1, f.9. 1995. (Type PE!).
泸定蛾眉蕨 lu ding e mei jue
Rhizomes stout, erect, densely clothed with brown, membranous, ovato-lanceolate scales at apices and stipe bases. Fronds caespitose; fertile fronds up to 97 cm; stipes 27 cm long, 5 mm thick at base; laminae oblong-lanceolate, ca. 70 cm long, ca. 21 cm broad, acuminate at apex, a little narrowed at base, sterile, pinnate- pinnatipartite; pinnae 26 pairs, sessile, alternate, narrowly deltoid-ovate to ovato-lanceolate, 5--11 cm long, 2--2.5 cm broad at middle, long-acuminate at apex, lower pinnae prominently broadened at base, pinnatipartite to the narrow wings of costae; segments 11--18 pairs, alternate, patent, oblong, 12--16 mm long, 3--5 mm broad, densely close each other, acute or obtuse-rounded at apex, toothed at margin, basal segments of pinna curved to raches. Veins visible, pinnate in segments, lateral veins 6--9 pairs, alternate, often forked. Laminae herbaceous when dried, green. Sori oblong to short-linear; indusia oblong or lunate, membranous, pale-brown, erose at margin, persistent. Spores bilateral, perispore surface with more dense rugate-saccate or papillate processes.
* About 3600 m. Sichuan (Luding).
This species is close to Lunathyrium wilsoni, but different in the lower pinnae ovato-triangular or ovato-lanceolate, prominently enlarged at base, pinna base more or less curved to the raches, segments acute or obtuse-rounded at apex, dentate at margin.
13. Lunathyrium wilsoni (Christ) Ching in Acta Phytotax. Sin. 9:71. 1964.
峨山蛾眉蕨 e shan e mei jue
Athyrium wilsoni Christ in Bull. Herb. Boiss. ser. 2. 3:512. 1903.
13a. var. wilsoni
峨山蛾眉蕨 e shan e mei jue
Rhizomes stout, erect, densely scaly at apices and stipe bases, scales brown, membranous, broadly lanceolate to linear-lanceolate ca. 1.5 cm long, 4 mm broad. Fronds caespitose; fertile fronds up to 1--1.5 m; stipes (10--) 20--25(--47) cm long, (2--)3--4(--6) mm thick, bases scaly, more sparse upwards; laminae oblong to narrowly oblong, (25--)60--90(--100) cm long,(10--)15--24(--28) cm broad, pinnatifid-acuminate at apex, pinnate, pinnae pinnatipartite; pinnae 15--20(--30) pairs, only 2--3 pairs of lower pinnae a little shortened, the basal pair ca. 7 cm, not auriculate, ca. 3.5 cm distant, subopposite, upward alternate, middle pinnae lanceolate or narrowly lanceolate, (5--)12--15(--20) cm long, (1--)2--3 cm broad, long-acuminate at apex, bases broader, subtruncate or broadly cuneate, pinnate parted to the wings of costae; segments (10--)20--25 pairs, oblong, 5--7(--10) mm long, 3--5 mm broad, close or spaced, roundish or near truncate at apex, sparsely crenulate or subentire at margin. Veins visible, pinnate in segments, lateral veins 6--8 pairs, simple. Laminae herbaceous or thick herbaceous when dried, green, sparsely bearing thin, short multicellular hairs on the surfaces along raches, costae and veins. Sori oblong to short-linear, 1.5--2 mm, 3--5(--6) pairs per segment (up to 6 pairs in the basal segments); indusia same with sori in shape, occasionally J-shaped or diplazioidal on the top of lamina and pinnae, subentire at margin, pale-brown. Spores bilateral, perispore surface with more dense rugate-saccate or papillate processes. 2n=80@.
* Damp areas, under forest on mountain slopes or streamside; 1400--3700 m. S Gansu, W Hubei, W Sichuan, Guizhou, NW Yunnan, SE Xizang. Type from Hubei (Changyang).
13b. var. incisoserratum Ching et Z. R. Wang in Acta Phytotax. Sin. 32(1):92. 1994 (Holotype PE!).
锐裂蛾眉蕨 rui lie e mei jue
Lunathyrium incisoserratum Ching in Y. L. Zhang et al., Sporae Pterid. Sin. 232. 1976, nom. nud., quoad nom., excl. tab.
It is very close to, but differs from the typical variety in segments usually bigger, narrowly triangular, distant, prominently toothed at margin, lateral veins sometimes 2-forked.
* Streamside under forests or shrub groves, wet areas in grass; 2800--3500 m. NW Yunnan (Deqen).
13c. var. habaense Ching et Z. R. Wang in Acta Phytotax. Sin. 32(1):91--92.1994 (Holotype PE!).
哈巴蛾眉蕨 ha ba e mei jue
It differs from the typical variety in segments narrowly triangular, separated each other by broad space, with long teeth or shallowly lobed at margin, lateral veins frequently forked.
* In valley under forests; ca. 2800 m. NW Yunnan (Zhongdian).
13d. var. maximum Ching et Z. R. Wang in Acta Phytotax. Sin. 32:92. 1994 (Holotype PE!).
大蛾眉蕨 da e mei jue
Lunathyrium maximum Ching in Y. L. Zhang et al., Sporae Pterid. Sin. 233, t. 46, f. 4--5. 1976. nom. nud.
It differs from the typical variety in plant larger, pinnae and segments broader, segments prominently toothed at margin.
* Under mixed forests, wet area; 2200--3100 m. S Gansu and Sichuan. Type from Sichuan (Li Xian).
13e. var. muliense Z. R. Wang in Acta Phytotax. Sin. 32(1):92. 1994 (Holotype PE!).
木里蛾眉蕨 mu li e mei jue
It differs from the typical variety in raches and costae on the abaxial surface hairs more thick and long, segments toothed at margin.
* Under coniferous forests on mountain slopes; 3300--3800 m. Sichuan, Xizang. Type from Sichuan (Muli).
14. Lunathyrium vermiforme Ching, Boufford et Shing in Journ. Arnold Arbor. 64:23, f. 7, a--c. 1983. (Holotype PE! Isotypes: A, CM, HIB, KUN, KYO, NA, NAS, NY, SFDH, UC, WH)
湖北蛾眉蕨 hu bei e mei jue
Rhizomes short, erect, clothed with dark-brown, membranous, lanceolate scales at apices and stipe bases. Fronds caespitose; fertile fronds 74 cm; stipes less than 20 cm long, ca. 2 mm thick, bases clothed with more dense scales, upward dark-stramineous, gradually glabrate; laminae oblong-lanceolate, ca. 55 cm long, ca. 18 cm broad at middle, acuminate at apex, gradually narrowed toward base, pinnate-pinnatipartite; pinnae ca. 20 pairs, patent, separated each other by narrow space, numerous pairs of lower pinnae gradually shortened towards the base, space between pinnae broader, the basal pair auriculate, ca. 1 cm, middle pinnae lanceolate to narrowly lanceolate, somewhat falcate, curved up, ca. 10 cm long, ca. 1.6 cm broad at base, acuminate at apex, subtruncate at base, pinnatipartite at margin; segments ca. 20 pairs, close, oblong, ca. 7 mm long, ca. 4 mm broad, subentire at margin, obliquely truncate or obtuse-rounded at apex. Veins visible on surfaces, pinnate in segments, lateral veins 4--5 pairs, simple. Laminae herbaceous when dried, green, pinnae subglabrous on surfaces, raches and costae on the abaxial surface sparsely bearing thin, short multicellular hairs. Sori short-linear, never curved even on the top segments of lamina or pinnae, 3--4 pairs per segment; indusia same with sori in shape, subentire. Spores bilateral, surface with rugate, rugate-saccate or papillate processes.
* In valley, damp area; ca. 1500 m. W Hubei (Lichuan).
15. Lunathyrium shennongense Ching, Boufford et Shing in Journ. Arnold Arbor. 64:21. 1983. (Holotype PE! Isotypes: A, CM, HIB, KUN, KYO, NA,NAS, NY, SFDH, UC, WH).
华中蛾眉蕨 hua zhong e mei jue
Lunathyrium centrochinense ("centro-chinense, centrochinensis") Ching ex K. H. Shing.
Rhizomes stout, erect or ascending, clothed with brown or blackish brown, membranous, broadly lanceolate, big scales at apices and stipe bases. Fronds caespitose; fertile fronds (30--)70(--100) cm; stipes (8--)12(--17) cm long, 2(--4) mm thick, stramineous or brown-reddish, grooved adaxially, sparsely bearing thin, short hairs or subglabrous; laminae oblanceolate, or oblong- oblanceolate, (25--)60--80 cm long, (6--)15--20 cm broad, acuminate at apex, gradually narrowed toward base, pinnate-pinnatipartite; pinnae(15--)20--22(--30) pairs, 6--10 pairs of pinnae gradually shortened towards the base, basal pair of pinnae often shortened as deltoid small auricle only ca. 1 cm, subopposite, apart from the second pair of pinnae more widely, middle pinnae narrowly lanceolate, (3--)10--12 cm long, the broadest at the bases, 1--2 cm broad, long-acuminate at apex, bases near truncate, sessile, alternate, 1--3 cm apart from each other, patent or slightly upswept, pinnatipartite; segments ca. 22 pairs, oblong, the middles (3--)7--8 mm long, 3(--5) mm broad at base, the basal pair a little longer, rounded-obtuse and crenate at apex, subentire or crenate at margin. Veins visible on surfaces, pinnate in segments, lateral veins 5--7 pairs per segment, simple. Laminae herbaceous when dried, green, raches and costae on the abaxial surface sparsely bearing short multicellular hairs or nearly glabrous, on the adaxial surface sparsely bearing brown short hairs. Sori elliptic or short-linear, 1.5--2 mm, (3--)4--5(--6) pairs per segment; indusia same with sori in shape, occasionally J-shaped on the apices of lamina and pinnae, pale-brown, slightly erose or subentire at margin. Spores bilateral, perispore surface with rugate, rugate-saccate or papillate processes. 2n=80.
* Mountain slope, under forests, wet places; 200--3300 m. Shaanxi, Anhui, Zhejiang, Jiangxi, Henan, Hubei, Hunan, Sichuan, Guizhou, Yunnan. Type from Hubei (Shennongjia).
This species is similar to Lunathyrium dolosum in shape, but hairs thin, short, sparse or subglabrous on the abaxial surface of raches and costae. It is a diploid, while the later is a tetraploid.
16. Lunathyrium truncatum Ching ex Z. R. Wang in Fl. Reip. Pop. Sin. 3(2):510-511, 1999. (Type PE!).
截头蛾眉蕨 jie tou e mei jue
Rhizomes ascending, clothed with brown, membranous, broadly lanceolate scales at apices and stipe bases. Fronds caespitose; fertile fronds 50--60 cm; stipes ca. 10 cm long, ca. 2 mm thick, stramineous, slightly bearing multicellular hairs and narrowly scales; laminae oblong, 40--50 cm long, 14 cm broad at middle, acuminate at apex, pinnate, pinnae pinnatipartite; pinnae ca. 20 pairs, 5--6 pairs of pinnae gradually shortened, ca. 5 mm, middle pinnae 8--9 cm long, 1.5 cm broad, linear-lanceolate, long-acuminate at apex, bases near truncate, alternate, 2.5--3 cm apart from each other, slightly upswept, pinnatilobed up to 2/3; segments ca. 15 pairs, close, almost as broad as long or a little longer, ca. 4 mm, square or oblong, truncate or retuse at apex, entire or subentire at margin. Veins visible abaxially, ca. 5 pairs per segment, simple. Laminae papery when dried, greenish-brown, raches stramineous, sparsely bearing multicellular long hairs, costae sparsely bearing short multicellular hairs abaxially. Sori short-linear, 1.5--2 mm, 2--3 pairs per segment, close each other; indusia same with sori in shape, entire or slightly erose at margin, yellow-brown, persistent. Spores bilateral, perispore surface with more dense papillate processes.
* About 3500 m. NW Yunnan.
17. Lunathyrium brevipinnum Ching et K. H. Shing ex Z. R. Wang in Acta Phytotax. Sin. 32(1):85--86, t. 2, f. 5--6. 1994 (Holotype PE! Isotypes KUN, PYU!).
短羽蛾眉蕨 duan yu e mei jue
Lunathyrium latibasis Ching, 1976, non Ching 1983.
Rhizomes short, erect, scaly at apices and stipe bases; scales brown, membranous, lanceolate, ca. 4 mm. Fronds caespitose; fertile fronds ca. 30 cm; stipes 5--7 cm long, rarely longer, ca. 1.4 mm thick, stramineous, subglabrous; laminae lanceolate, 20--25 cm long, ca. 6 cm broad at middle, acuminate at apex, slightly narrowed toward base, pinnate-pinnatipartite; pinnae ca. 15 pairs, patent, sessile, the uppers alternate, the lowers subopposite, ca. 2 cm apart from each other, the lower 2 pairs a little shortened, the basal pair ca. 1.5 cm, not auriform, middle pinnae lanceolate, ca. 3 cm long, broader, 1--1.3 cm broad at base, acute or short-acuminate at apex, pinnatipartite to the wings of costae; segments 6--8 pairs, close, deltoid-oblong, obtuse, rarely obtuse-acute at apex, entire at margin, upswept, the basal pair a little bigger, up to 5 mm long, ca. 2.6 mm broad, sparsely crenate. Veins visible abaxially, pinnate in segments, lateral veins 3--4 pairs, up to 5 pairs in the basal acroscopic segments, simple, upswept. Laminae herbaceous when dried, greenish brown, subglabrous, only costae sparsely with short, brown, thin multicellular hairs abaxially. Sori small, oblong-ovate, 3--6 per segment, upswept; indusia short-lunate, thick, slightly erose at margin, pale-green, persistent. Spores bilateral, perispore surface with denser rugate-saccate or papillate processes.
* In shrub groves; 2200--3000 m. Yunnan. Type from Yunnan (Dayao).
This species is close to Lunathyrium shennongense in shape, but smaller, pinnae fewer, usually less than 12 pairs, shorter, usually ca. 3 cm, no more than 5 cm, a little broadened at bases, segments of pinnae fewer, only 5--7 pairs, segments rounded-obtuse or subtruncate at apex, so it is easy to be recognized.
18. Lunathyrium dolosum (Christ) Ching in Acta Phytotax. Sin. 9:72. 1964.
昆明蛾眉蕨 kun ming e mei jue
Athyrium dolosum Christ, Bull. Acad. Geogr. Bot. Mans 136. 1907 (Syntype P!); Deparia dolosa Kato in sched.
18a. var. dolosum
昆明蛾眉蕨 kun ming e mei jue
Rhizomes erect or ascending, clothed with brown-red, membranous, broadly lanceolate big scales at apices and stipe bases. Fronds caespitose; fertile fronds (30--)100--120 cm; stipes (6--)24 cm, stramineous or slightly brown-reddish, bases clothed with brown, ovato-lanceolate scales, upward clothed with thin scales or tortuous, hyaline, multicellular hairs; laminae narrowly oblong-lanceolate or narrowly oblanceolate, (25--)80 cm long, (10--)15(--25) cm broad, pinnatifid-short-acuminate at apex, gradually narrowed toward base, pinnate-pinnatipartite; pinnae (10--)30 pairs or more, the lowers subopposite, upward alternate, patent or slightly upswept, ca. 2 cm apart from each other, 7--8 pairs of lower pinnae gradually shortened towards the base, the basal pair only 1--2 cm, broader, triangular, middle pinnae lanceolate to narrowly lanceolate, (4--)8--10(--13) cm long, bases broader, 1--2 cm broad, acuminate at apex, bases subtruncate, or broadly cuneate, pinnatipartite; segments (10--)15(--20) pairs, oblong, (3--)8--9(--12) mm long, 4--5 mm broad, a little ascendant, obtuse or obtuse-rounded at apex, joining each other by broad wing at base, toothed at margin, separated each other by narrow space. Veins visible on surfaces, pinnate in segments, lateral veins 4--5(--7) pairs, simple, sometimes forked in the basal segments. Laminae herbaceous when dried, brown-green, raches, costae and costules and veinlets sparsely bearing longer multicellular hairs on surfaces. Sori oblong or short-linear, in some degree J-shaped at pinna top, 1.5--2 mm, 3--5 pairs per segment; indusia same with sori in shape, brownish, slightly erose at margin, persistent. spores bilateral, perispore surface with rugate, rugate-saccate or papillate processes. 2n=160.
* In valley, under forests, wet area; 1900--3500 m. SW Sichuan and Yunnan. Type from Yunnan (Kunming).
18b. var. chinense Z. R. Wang in Fl. Reipubl. Popul. Sin. 3(2):316. 1999. (Type PE!).
中华蛾眉蕨 zhong hua e mei jue
Lunathyrium chinense Ching in Y. L. Zhang et al., Sporae Pterid. Sin. 231. 1976. nom. nud., quoad nom, excl. tab.
It differs from the typical variety in segments deltoid-oblong, subentire at margin.
* Under forests; ca. 2300 m. SW Yunnan (Gengma).
19. Lunathyrium orientale Z. R. Wang et J. J. Chien ex J. J. Chien in Acta Phytotax. 22:228, t. 1, f. 2. 1984 (Holotype PE!).
东亚蛾眉蕨 dong ya e mei jue
Deparia pycnosora var. albosquamata Kato (Holotype TI! Isotype KYO, TNS, PE!); Deparia albosquamata Nakaike; Athyrium pycnosorum Christ, pro parte; Lunathyrium pycnosorum Koidz. pro parte.
19a. var. orientale
东亚蛾眉蕨 dong ya e mei jue
Rhizomes erect or ascending, densely clothed with brown or dark-brown, membranous, broadly lanceolate scales at apices and stipe bases. Fronds caespitose; fertile fronds (30--)70--80 cm; stipes (5--)10--14(--20) cm long, (2--)3--4 mm thick, stramineous or red-brownish, clothed with rather dense, zigzag, thick multicellular hairs and thin scales on their lower parts, upwards gradually sparse; laminae oblanceolate or oblong-lanceolate, (25--)50--60(--65) cm long, (8--)12--15(--20) cm broad, pinnatifid and short-acuminate at apex, gradually narrowed toward base, pinnate-pinnatipartite; pinnae (15--)--20 pairs, numerous pairs of lower pinnae gradually shortened towards the base, the basal pair often only 0.5--1 cm, subopposite, upward alternate, middle pinnae narrowly lanceolate, (4--)8--10 cm, bases broader, 1--2 cm broad, long-acuminate at apex, bases near truncate, patent or slightly upswept; segments ca. 12 pairs, oblong, (3--)5--8 cm long, 3--4(--5) mm broad, close each other, obtuse-rounded at apex, subentire at margin, in some cases crenulate. Veins visible on surfaces, pinnate in segments, lateral veins 5--7 pairs, simple. Laminae herbaceous when dried, raches and costae bearing thick multicellular hairs abaxially. Sori oblong or lunate, sometimes J-shaped at apices of lamina and pinnae, 1.5--2 mm, 3--5 pairs per segment; indusia same with sori in shape, pale-brown, slightly erose or shortly ciliated at margin, persistent; spores bilateral, perispore surface with rather densely rugose-saccate or papillate processes.
Under forests, wet places; 900--2200 m. Taiwan, Anhui [CS Japan]. Type from Japan (PE!).
19b. var. jiulungense (Ching) Z. R. Wang in Fl. Reip. Pop. Sin. 3(2):318, 1999. (Type PE!).
九龙蛾眉蕨 jiu long e mei jue
Lunathyrium jiulungense Ching in Bull. Bot. Res. 2(2):71--72, pl. 3. f. 3. 1982; Lunathyrium orientale var. huangshanense Z. R. Wang (Holotype PE!).
It differs from the typical variety in indusia shortly ciliated at margin, stipes stramineous, multicellular hairs straight on stipes and raches.
* In rock crevice or on mountain slope, under forests, wet area; 900--2700 m. Anhui, Zhejiang, Jiangxi, Sichuan. Type from Zhejiang (Jiulong Shan, Suichang).
From abortive spores and some morphological features many hybrids can be estimated from the specimens. Here only four hybrids have been published as species are listed.
Lunathyrium Χ kanghsienense Ching et Y. P. Hsu (pro sp.) in Fl. Tsinling. 2:94,212--213, 1974. (Type PE!)
康县蛾眉蕨 kang xian e mei jue
The type of this hybrid morphologically is in between of L. liangshanense var. sericeum and L. sichuanense, similar to the former in indusia with multicellular long hairs on the back. But, segments bearing no multicellular long hairs at margin, at most occasionally with a few short hairs between teeth. The hairs on the surfaces of segments are much more than the latter. Spores mostly abortive.
* 1300--1400 m. Gansu (Kangxian).
Lunathyrium Χ nanchuanense Ching et Z. Y. Liu (pro sp.) in Bull. Bot. Res. 3(4):15--16, t. 14. 1983. (Type PE!)
南川蛾眉蕨 nan chuan e mei jue
This hybrid is morphologically in between of L. shennongense and L. vegetius. The hairs on the abaxial surface of raches and costae are very thin, short and sparse, multicellular hairs absent at the incision between segments, the multicellular hairs on the adaxial surface of veins of fertile pinnae very short, that is same to L. vegetius. But lower pinnae gradually shortened, spore surface with rugate folds or baculate processes, that is same to L. shennongense. Spores partly abortive in a sporangium.
* Under forests; 1600--1700 m. Chongqing (Jinfo Shan, Nanchuan).
Lunathyrium Χ subimbricatum (Ching) Z. R. Wang (Type PE!).
栉比蛾眉蕨 jie bi e mei jue
Lunathyrium subimbricatum Ching in Y. L. Zhang et al., Sporae Pterid. Sin. 233--234, t. 45, f. 21,25. 1976. nom. nud. fig. mala.
The hybrid is morphologically similar to L. sichuanense, but plant is far bigger, basal segments of pinna prominently broadened. It looks also with relationship to L. wilsoni. Spores completely abortive.
* Under broad leaf forests on mountain slopes, rock slits waterside; 2700--2800 m. W Yunnan (Yongping).
Lunathyrium Χ heterocarpum (Ching) Z. R. Wang (Type PE!)
杂果蛾眉蕨 za guo e mei jue
Lunathyrium heterocarpum Ching, in Y. L. Zhang et al., Sporae Pterid. Sin. 232, t. 46, f. 10,15. 1976. nom nud. fig. mala. (Type PE!)
This hybrid is morphologically in between of L. dolosum and L. shennongense, pinnae up to 30 pairs or more, segments up to 20 pairs or more, raches and costae on the abaxial surface bearing rather dense multicellular long hairs. Spores partly abortive, and the good spores are great different in size in a same sporangium.
* Under forests on mountain slopes; ca. 3300 m. Sichuan (Li Xian).
ATHYRIUM Roth in Rom. Mag. 2(1):105. 1799.
蹄盖蕨属 ti gai jue shu
Terrestrial, medium-sized herbal plants. Rhizomes short, mostly erect, occasionally creeping or ascending, rarely long-creeping; fronds caespitose, rarely approximate or distant. Stipes long, often swollen at bases, protuberant adaxially and depressed abaxially, laterally with uniserial tuberculate pneumathodes, sharpened towards the base, in some cases bearing no pneumathodes, not sharpened towards the base; with 2 vascular bundles in section, upward to raches conjoint as U-shaped; stipe scaly at base, scales red-brown, brown or blackish brown, ovato-lanceolate, linear-subulate or linear-lanceolate, entire, consist of linear wall-thickened cells, membranous, basifixed; stipes grooved adaxially, upward open to raches, bearing short glandular-hairs in the grooves; laminae ovate, oblong or broadly lanceolate, pinnate to 3-pinnate, pinnae, pinnules or second pinnules connate or separate with raches, costae or costules at base, if connate usually decurrent at base， raches, costae or costules protuberant abaxially, grooved adaxially, the grooves of rachis and costae are open to the grooves of costae and costules, a fleshy spine-like process present or absent in adaxial groove margin at costa base. Veins free, forked or pinnate, veinlets extend to the tooth tip. Laminae usually herbaceous when dried, in a few cases papery or leathery, usually glabrous, rarely bearing scales or hairs, only on the adaxial surface of raches, costae or costules frequently bearing short glandular-hairs. Sori round, round-reniform, horseshoe-shaped, J-shaped, oblong or short-linear, dorsal, lateral or cross over veinlets; indusia round-reniform, horseshoe-shaped, J-shaped, lunate, oblong or short-linear, brown, membranous, erose or ciliated, rarely entire at margin, usually persistent, rarely exindusiate or indusia undeveloped. Spores bilateral, elliptic in polar view, bean-shaped in equatorial view, and perispore surface with folds or without folds. X=40.
About 160 species in the world, occur mainly in the temperate zone and under subtropical mountain forests. About 117 species with a number of varieties and hybrids known in China, occur in all provinces and districts. The distribution center is the mountain area of SE China.
ATHYRIUM is one of the largest genera of pteridophytes and one of most difficult genera to treat taxonomically. Molecular data shows ATHYRIUIM is polyphyletic. It is the trunk of the ATHYRIACEAE. Hybridization makes the taxonomy of this genus more difficult. There are more than 60 hybrids have been known in Japan. Much more hybrids have not been studied in China.
About 300 "species" names of ATHYRIUM from China were published. They are treated as 117 species with a number of varieties and hybrids.
Key to the species
1a. Sori rotund or elliptic; exindusiate or indusia caduceus; spore perispore with folds.
2a. Stipes and raches dark brown, bearing throughout dense castaneous black or dark brown, ovato-lanceolate scales and mixed with fibrous ones; basal pinnae tapering towards the base; pinna apex obtuse; stipes far shorter than laminae. Northwest Yunnan, West Sichuan and Southeast Xizang
1. A. wallichianum
2b. Stipes and raches stramineous or pale purplish red, only at the base bearing many brown scales; basal pinnae only 1--2 pairs a little abbreviated; stipes a little shorter than laminae; pinna apex acuminate.
3a. Laminae pinnate, pinnae pinnatilobate to pinnatipartite, occasionally 2-pinnate; pinnae sessile or subsessile (stipes less than 3 mm); teeth at lobe margin short and pointing outwards. Hunan, Guangxi, Sichuan, Guizhou and Yunnan
2. A. dissitifolium
3b. Laminae 2-pinnate; pinnae prominently stipitate (stipes up to 10 mm or more); teeth at lobe margin long and pointing forwards. NW Yunnan and Xizang
3. A. exindusiatum
1b. Sori elliptic, short-linear, J-formed, horseshoe-shaped or reniform; indusia persistent, rarely incomplete but seeable, at least when tender; spore perispore with or without folds.
4a. Rhizomes long- or short-creeping; fronds distant or approximate; or rhizome erect, but spore perispore with folds, pinnae sessile not auricled at acroscopic bases.
5a. Rhizomes long-creeping; costae and costules bearing prominent spine-like processes adaxially; spore perispore with or without folds.
6a. Middle plant, more than 20 cm high; laminae 3-pinnate or more divided; along costae and costules two sides with narrow wings; sori not close to costae or costules, indusia not overlay costae or costules; spore perispore with folds. Xizang
4. A. pectinatum
6b. Small plant, less than 20 cm high; laminae pinnate, pinnae pinnatipartite; sori close to costae or costules, indusia often overlay costae or costules; perispore without folds. Northwest Yunnan
100. A. chingianum
5a. Rhizomes creeping or erect, but pinnae sessile not auricled at acroscopic bases; costae and costules without processes adaxially; spore perispore with folds.
7a. Rhizomes long-creeping; fronds distant.
8a. Laminae subdimorphic, stipes of sterile frond shorter than laminae, but the one of fertile frond far longer than laminae; laminae deltoid-ovate, pinnae 3--5 pairs; basal pair of pinnae not abbreviated, basiscopic lobes of costa longer than acroscopic ones. Yunnan
5. A. ruilicola
8b. Laminae monomorphic, stipes shorter than laminae or equal long; laminae ovato-oblong; pinnae 15--20 pairs; basal pair of pinnae abbreviated, basiscopic lobes of costa equal to acroscopic ones.
9a. Laminae 2-pinnate, supra-surface glabrous; teeth not long-apiculate. Yunnan, Sichuan and Xizang
6. A. biserrulatum
9b. Laminae 3-pinnate, adaxial surface pubescent; teeth long-apiculate. Yunnan
7. A. wumonshanenicum
7b. Rhizomes short-creeping (occasionally long-creeping), fronds approximate, or rhizome erect, but pinnae sessile not auricled at acroscopic bases.
10a. Laminae ovate to narrowly ovate, apex cuspidate, basal pinnae only 1--2 pairs a little shortened; pinnae stipitate; stipes only a little shorter than laminae; scales at stipe bases pale brown; sori short-linear, oblong or long J-formed.
11a. Pinnae usually pinnate; pinnules ovate, oblong-lanceolate or falciform-lanceolate, shallowly lobed to pinnatipartite at margin, basal pinnules usually shortly stipitate. Liaoning, Nei Mongol, Beijing, Hebei, Shanxi, Henan, Shandong, Shaanxi, Gansu, Ningxia, Hubei, Hunan, Anhui, Jiangxi, Zhejiang, Jiangsu, Guangxi, Guangdong, Sichuan, Guizhou, Yunnan and Taiwan
8. A. niponicum
11b. Pinnae usually pinnatipartite to pinnatisect; lobes falciform-lanceolate, at most shallowly lobed at margin, decurrent and connate with the broad wings of costa at the basiscopic base. Yunnan
9. A. brevisorum
10b. Laminae lanceolate, oblong-lanceolate or oblanceolate, apex acuminate; pinnae sessile or occasionally shortly stipitate (stipe no more than 5 mm); stipes far shorter than laminae; scales at stipe bases red-brown or castaneous; sori subrotund, elliptic, reniform or short J-formed.
12a. Indusia pale brown, entire, basal pinnae not shortened or almost not shortened; rhizome erect, pinnae sessile not auricled at acroscopic bases.
13a. Scales at stipe bases yellow-brown, laminae pinnate-pinna pinnatipartite to 2-pinnate-pinnule pinnatilobate, pinnae narrow, lanceolate. Heilongjiang, Jilin, Liaoning, Shandong, Anhui, Jiangsu, Zhejiang, Jiangxi, Hunan, Sichuan and Guizhou
10. A. yokoscense
13b. Scales at stipe bases red-brown, black-brown at the middle, laminae pinnate, pinnae pinnatilobate to pinnatipartite, broad, ovate
11. A. fauriei
12b. Indusia pallid, margin ciliated; basal pinnae many pairs gradually shortened towards the base or only 1--2 pairs prominently shortened; rhizome short-creeping.
14a. Teeth at the lobe margin obtuse. Heilongjiang, Jilin, Liaoning, Nei Mongol, Beijing, Hebei, Shanxi, Nei Mongol, Henan, Hubei, Shaanxi, Gansu and Ningxia
12. A. fallaciosum
14b. Teeth at the lobe margin sharp and long.
15a. Raches and costae glabrous abaxially. Sichuan, Yunnan and Xizang
13. A. rupicola
15b. Raches and costae pubescent abaxially. Sichuan and Yunnan
14. A. glandulosum
4b. Rhizomes erect or ascendant, fronds caespitose.
16a. Costae (or costules and midribs) without spine-like processes adaxially, at most on the costa top of bigger plants rarely with extremely short processes, spore perispore with or without folds.
17a. Pinnae prominently broadened at base, asymmetrical, auricled at acroscopic bases, usually cuneate at basiscopic bases; when laminae pinnate or more divided, all lobes or pinnules anadromic; costae without spine-like processes on the adaxial surface; spore perispore with folds (rarely without folds).
18a. Small plants, fertile fronds less than 10 cm, spore perispore without folds. Taiwan
15. A. minimum
18b. Middle plants, fertile fronds more than 10 cm, spore perispore with folds.
19a. Laminae pinnate, pinnae margin repand or shallowly deltoid-lobed, occasionally acroscopic basal lobes of basal pinnae pinnatisect; raches glandular pubescent.
20a. Pinna obtuse at apex, basal acroscopic auricled lobes entire or slightly repand to dentate, raches with sparse black or brown glandular hairs. Taiwan, Yunnan and Xizang
16. A. nakanoi
20b. Pinna acute at apex, bases acroscopic auricled lobes denticulate at apex, raches with sparse brown glandular hairs. West Guizhou
17. A. zhenfengense
19b. Laminae pinnate to 2-pinnate or more divided, if pinnate, lobes of pinnae not deltoid; raches without glandular hairs (occasionally with pale, brown, short unicellular hairs).
21a. Laminae ovate to ovate, basal pair of pinnae biggest, long ovate, basal 1--2 pairs pinnules of basal pinnae a little shortened; ultimate pinnules or lobes obtuse. Yunnan and Xizang
18. A. bucahwangense
21b. Laminae narrowly lanceolate to ovato-lanceolate, basal pair of pinnae a little shortened, deltoid-ovate to deltoid-lanceolate, basal pair of pinnules of basal pinnae biggest; ultimate pinnules or lobes acute or rotund.
22a. Raches without glandular hairs adaxially. Taiwan
19. A. erythropodum
22b. Raches glandular pubescent adaxially.
23a. Laminae thickly papyraceous or thin-coriaceous, dark green, surface splendent; indusia small, partly fugacious when sori ripe. Sichuan, Guizhou, Yunnan, Xizang and Taiwan
20. A. drepanopterum
23b. Laminae papyraceous or herbaceous, green, surface not splendent; indusia bigger, persistent.
24a. Laminae 2-pinnate to 3-pinnate; stipes pale purplish red, scales at bases reddish brown; pinnae and ultimate pinnules acute at apex; sori less than 1 mm in diameter. Hunan, Sichuan, Yunnan and Xizang
21. A. fimbriatum
24b. Laminae pinnate to 2-pinnate; stipes stramineous, scales at bases brown or dark brown; pinnae and ultimate lobes rotund at apex; sori 1--2 mm in diameter.
25a. Laminae usually lanceolate, pinnate, at most 2-pinnate at bases, but separate pinnules no more than 5 pairs. Hunan, Jiangxi, Taiwan, Guangdong, Guangxi, Sichuan, Guizhou, Yunnan and Xizang
22. A. anisopterum
25b. Laminae usually ovato-lanceolate, 2-pinnate, separate pinnules of middle pinnae may more than 10 pairs. Taiwan, Yunnan and Xizang
23. A. foliolosum
17b. Pinnae narrowed at bases, not or a little broadened, symmetric or subsymmetric; at least pinnules or lobes of supra-medial pinnae catadromic or opposite; costae of bigger plants rarely with very short processes adaxially; spore perispore without folds.
26a. Laminae densely pubescent on adaxial surface.
27a. Raches and costae pubescent abaxially; lobes of pinnae obtuse at apex. Sichuan, Jinfo Shan
24. A. sericellum
27b. Raches and costae glabrous abaxially; lobes of pinnae rotund at apex. Sichuan, Ebian
25. A. suprapubescens
26b. Laminae glabrous.
28a. Basal pinnae prominently constricted at bases; sori mostly short-linear; raches and costae glabrous or with short straight glandular hairs abaxially.
29a. Laminae deltoid-ovate, apex cuspidate, basal several pairs of pinnae subopposite, basiscopic lobes of basal pair of pinnae longer than acroscopic ones. Hainan
26. A. hainanense
29b. Laminae lanceolate or ovato-oblong, apex acuminate, pinnae alternate, basiscopic lobes of basal pair of pinnae almost equal to acroscopic ones.
30a. Indusia incomplete, only seeable when tender, teeth on lobe margin cuspidate. Guangxi, Guizhou and Yunnan
27. A. pachyphyllum
30b. Indusia complete, persistent, teeth on lobe margin not cuspidate. Yunnan
28. A. wangii
28b. Basal pinnae a little broadened or slightly constricted at bases; sori mostly reniform, horseshoe-shaped, J-formed or oblong; raches and costae sparsely bearing squarrose-glandular hairs abaxially.
31a. Laminae broadly ovate or ovate, basal 1--2 pairs of pinnae a little shortened or almost not shortened. Northeast.
32a. Scales at stipe bases black, splendent. Heilongjiang and Jilin
29. A. melanolepis
32b. Scales at stipe bases brown or dark brown, not splendent.
33a. Plant strong, laminae herbaceous, stipes and raches stramineous or pale purplish red, surface not splendent. Heilongjiang, Jilin, Liaoning, Nei Mongol, Beijing, Hebei, Shanxi and North Shandong
30. A. brevifrons
33b. Plant thin and smaller, laminae thin herbaceous, stipes and raches pale reddish, surface splendent. Northeast Heilongjiang
31. A. rubripes
31b. Laminae narrowly ovate, elliptic-lanceolate or oblanceolate, basal pinnae more than 2--3 pairs gradually shortened. Occur from the North to the Southwest.
34a. Laminae narrowly ovate, basal 2--3 pairs of pinnae shortened, but the basal pair of pinnae not shortened as auriculate. Hebei, Shanxi, Nei Mongol, Shandong, Henan, Shaanxi, Gansu and Ningxia
32. A. sinense
34b. Laminae elliptic-lanceolate or oblanceolate, basal pinnae more than 4 pairs gradually shortened, the basal pair of pinnae often shortened as auriculate. Occur from S Qinling to Xizang.
35a. Scales at stipe bases lanceolate or linear-lanceolate, less than 1.5 mm wide, laminae pinnate-pinnae pinnatipartite, lobe margin serrate, lateral veinlets usually simple. Sichuan and Yunnan
33. A. attenuatum
35b. Scales at stipe bases ovate or deltoid-ovate, more than 3--5 mm wide, laminae usually 2-pinnate, pinnules margin shallowly lobed or biserrate, lateral veinlets usually 2--4 furcated.
36a. Laminae black when dried; scales at stipe bases castaneous at middle, brown at margin; pinnule rounded-truncate at apex. NW Yunnan
34. A. tarulakaense
36b. Laminae not black when dried; scales at stipe bases brown or dark brown; pinnule apex obtuse.
37a. Pinnae deltoid-ovate or deltoid-lanceolate, less than 3--4 cm, approximate each other. Sichuan and Xizang
35. A. contigens
37b. Pinnae linear-lanceolate, more than 4 cm, separate by space from each other.
38a. Lateral pinnae shortly stipitate, slightly constricted towards bases, pinnules longer, may reach 2.5--3 cm. Yunnan
36. A. nudifrons
38b. Lateral pinnae subsessile, not constricted toward bases, pinnules usually less than 2.5--3 cm.
39a. Middle pinnae about 1 cm wide, separated from each other by space more than pinna wide. Gansu, Hubei, Sichuan and Yunnan
37. A. nephrodioides
39b. Middle pinnae more than 1 cm wide, separated from each other by space far less than pinna wide.
40a. Pinnae shorter, narrower, 3--6 cm long, less than 1.8 cm wide, prominently sloping upwards patent. Sichuan
38. A. adscendens
40b. Pinnae longer, wider, patent or slightly ascendant. Gansu, Gansu, Sichuan, Guizhou, Yunnan and Xizang
39. A. dentigerum
16b. Costae (or with costules and midribs) with long or short spine-like processes on the adaxial surface; perispore without folds.
41a. Laminae pinnate, oblanceolate, pinnae pinnatilobate to pinnatipartite, many pairs of basal pinnae gradually shortened, basal pair of pinnae often shortened as auriculate. Yunnan and Xizang
40. A. mehrae
41b. Laminae pinnate to 3-pinnate, not oblanceolate, basal pinnae not shortened, or at most one to several pairs a little shortened, basal pair of pinnae never shortened as auriculate.
42a. Laminae narrowly lanceolate, pinnae more than 20--25 pairs.
43a. Laminae usually less than 35 cm, pinnate to 2-pinnate, lobe margin dentate, costae with short spine-like processes adaxially. Zhejiang, Hunan, Jiangxi and Guizhou
41. A. multipinnum
43b. Laminae may reach more than 35 cm, 2-pinnate, lobe margin long-serrulate, costae with long pine-like processes adaxially. Jiangxi and Guangxi
42. A. elongatum
42b. Laminae broader, not narrowly lanceolate, pinnae less than 20--25 pairs.
44a. Indusia J-formed, horseshoe-shaped, reniform, elliptic, short-linear etc., lateral, crossing or dorsal on veinlets; scales at stipe bases often yellow-brown, brown or dark brown.
45a. Pinnules or lobes of supra-medial pinnae anadromic, occasionally catadromic or subopposite, raches and costae stramineous, rarely pale purplish red, glabrous or sparsely pubescent abaxially; the narrow wing margins of costae or the sinus between lobes glabrous.
46a. Pinnules or lobes of supra-medial pinnae opposite or subopposite.
47a. Pinnules or lobes downwards reflexed or subpatent. Taiwan
43. A. oppositipinnum
47b. Pinnules or lobes patent. Sichuan
44. A. crassipes
46b. Pinnules or lobes of supra-medial pinnae anadromic, alternate, or only basal pair subopposite.
48a. Pinnae (especially at laminae tip) or pinnules upwards patent or at most subpatent.
49a. Basal pinnae alternate, at most the basal pair subopposite; raches and costae pale purplish red, prominently with spine-like processes adaxially, but costules without. Taiwan
45. A. tripinnatum
49b. Bases pinnae one to several pairs opposite or subopposite; raches and costae stramineous, with very short spine-like processes adaxially, costules also with.
50a. The basal pair of pinnules subopposite, raches and costae sparsely pubescent abaxially. Fujian, Jiangxi, Hunan, Sichuan and Guizhou
46. A. deltoidofrons
50b. The basal pinnules anadromic, raches and costae glabrous abaxially. Guangxi, Sichuan, Guizhou, Yunnan and Xizang
47. A. delicatulum
47b. Pinnae (especially at laminae tip) or pinnules sloping downwards reflexed.
51a. Laminae pinnate, pinnae at most pinnatipartite, sessile, sloping downwards reflexed, except the one at laminae tip patent. Taiwan
48. A. reflexipinnum
51b. Laminae pinnate to 2-pinnate, pinnae shortly stipitate, usually patent or sloping upwards, only the ones at laminae tip downwards reflexed; pinnules may reach pinnatisect, sloping downwards or patent.
52a. Sori horseshoe-shaped, pinnules lanceolate, subsessile. Zhejiang, Fujian, Jiangxi, Guangxi, Sichuan, Guizhou, Yunnan and Xizang
49. A. devolii
52b. Sori oblong or J-formed, pinnules ovato-deltoid or oblong, prominently stipitate or the bases decursive and connected with the narrow wing of costa.
53a. Pinnules ovato-deltoid, acute at apex, prominently stipitate (stipes ca.1 mm). Zhejiang (Qingyuan)
50. A. baishanzuense
53b. Pinnules oblong, rotund-obtuse at apex, the bases decursive and connected with the narrow wing of costa. Zhejiang (Maoshan)
51. A. maoshanense
45b. Pinnules or lobes of supra-medial pinnae catadromic or subopposite; raches and costae usually pale purplish red, occasionally stramineous, pubescent abaxially; the narrow wing margins of costa or the sinus between lobes sparsely pubescent or glabrous.
54a. Indusia usually reniform, rotund-reniform or elliptic; laminae broadly ovate, the basal pair of pinnae biggest, apex acuminate; scales yellow-brown. Hubei, Hunan, Shaanxi, Gansu, Sichuan and Yunnan
52. A. omeiense
54b. Indusia usually oblong, J-formed or horseshoe-shaped; laminae usually oblong-ovate, apex cuspidate or acuminate, the basal pair of pinnae almost equal to the second pair or a little bigger; scales brown or dark brown.
55a. Pinnae prominently stipitate, stipes usually more than 2 mm, alternate, except the basal pair subopposite; the narrow wing margins of costa or the sinus between lobes glabrous, occasionally sparsely pubescent; indusia margin subentire or erose.
56a. The narrow wing margins of costae glabrous, raches and costae glabrous or sparsely pubescent.
57a. Pinnules oblong, rotund-obtuse at apex. Zhejiang and Anhui
53. A. intermixtum
57b. Pinnules ovate to oblong-lanceolate, acute.
58a. Lamina apex usually acuminate; raches and costae glabrous abaxially, pinnules of middle and basal pinnae pinnatipartite to pinnatisect.
59a. Scales pale brown, linear-lanceolate, costae pale purplish red abaxially. Sichuan
54. A. costulelisorum
59b. Scales dark brown, ovate to ovato-lanceolate, costae stramineous. Xizang
55. A. dentilobum
58b. Lamina apex often cuspidate; raches and costae sparsely pubescent abaxially, pinnules of middle and basal pinnae pinnatilobate to pinnatipartite.
60a. Costae pale purplish red; laminae thin herbaceous, apex cuspidate prominently. Henan, Anhui, Jiangxi, Zhejiang, Fujian, Taiwan, Hubei, Hunan, Guangxi, Sichuan, Guizhou, Yunnan, Shaanxi and Gansu
56. A. vidalii
60b. Costae stramineous; laminae papyraceous, apex cuspidate often not prominently. Hubei, Hunan, Guangxi, Sichuan, Guizhou, Yunnan, Shaanxi, Gansu and Xizang
57. A. mackinnonii
56b. The narrow wing margins of costae sparsely pubescent, costae densely pubescent abaxially.
61a. Laminae supra-surface and lobe margin sparsely pubescent, pinnules narrow-deltoid to deltoid-oblong, acute at apex. Sichuan
58. A. ludingense
61b. Laminae supra-surface and lobe margin glabrous, pinnules oblong to deltoid-ovate, obtuse at apex. Gansu and Sichuan
59. A. hirtiraches
55b. Pinnae usually sessile, occasionally shortly stipitate, stipes no more than 2 mm, basal 3 or more pairs often opposite or subopposite; the narrow wing margins of costa or the sinus between lobes usually sparsely pubescent, occasionally glabrous; indusia margin usually ciliated, occasionally lacerate or erose.
62a. Raches and costae stramineous, densely bearing unicellular straight hairs abaxially, with very short spine-like processes adaxially; but lobe midribs without. Sichuan, Yunnan and Xizang
60. A. himalaicum
62b. Raches and costae usually pale purplish red, occasionally stramineous; sparsely pubescent or glabrous abaxially, with long or short spine-like processes adaxially; costules or lobe midribs on with or without.
63a. Spine-like processes of costae on the adaxial surface very short, if long, costules or midribs without spine-like processes.
64a. Plant small, less than 30 cm high; pinnae less than 5 cm, pinnatipartite; raches and costae pale purplish red or stramineous; the narrow wing margins of costae sparsely pubescent.
65a. Basal pair of pinnae prominently shortened; spine-like processes of costae on the adaxial surface very short; raches and costae stramineous; laminae supra-surface glabrous. Sichuan
61. A. emeicola
65b. Basal pair of pinnae not shortened; spine-like processes of costae on the adaxial surface sharp and long; raches brown- stramineous, costae pale purplish red abaxially, laminae supra-surface sparsely bearing pallid short hairs. Xizang
62. A. bomicola
64b. Plant middle-small, more than 30 cm high; pinnae more than 7 cm, pinnate; raches and costae stramineous; the narrow wing margins of costae sparsely pubescent or glabrous.
66a. Spine-like processes of costae on the adaxial surface sharp and long; the narrow wing margins of costae glabrous. Xizang
63. A. flabellulatum
66b. Spine-like processes of costae on the adaxial surface very short; the narrow wing margins of costae sparsely with hairs.
67a. Middle-small plants, less than 50 cm high, pinnules oblong, about 7 mm, rounded at apex, margin crenate. Sichuan and Yunnan
64. A. daxianglingense
67b. Middle plants, may reach 1 m high, pinnules broadly lanceolate, 22--27 mm, obtuse, margin incised.
68a. Medial pinnae more than 20 cm; pinnules more than 20 pairs, oblong-lanceolate, or oblong-ovate, obtuse at apex. Yunnan
65. A. obtusilimbum
68b. Medial pinnae less than 12 cm; pinnules about 14 pairs, deltoid-ovate, or deltoid-broad-lanceolate, obtuse at apex. Sichuan
66. A. baoxingense
63b. Spine-like processes of costae on the adaxial surface longer, narrowly deltoid, broader at bases; if with pinnules, costules or midribs with spine-like processes too.
69a. The narrow wing margins of costae and raches glabrous.
70a. Laminae thin herbaceous, lobe teeth long and pointing directly outwards, sori 2--4 pairs on each pinnule. Yunnan
67. A. jinshajiangense
70b. Laminae papyraceous or hard herbaceous, lobe teeth short and not pointing directly outwards, sori 4--7 pairs on each pinnule. Yunnan
68. A. interjectum
69b. The narrow wing margins of costae and raches more or less with unicellular short hairs.
71a. Laminae supra-surface with unicellular short hairs. Sichuan and Yunnan
69. A. suprapuberulum
71b. Laminae adaxial surface glabrous.
72a. Pinnae and lobes small, pinnae less than 1 cm broad, lobes oblong, 2--3 mm wide, rounded-truncate at apex; scales at stipe bases black in centre, prominently with narrow pale brown margin. Xizang
70. A. zayuense
72b. Pinnae and lobes bigger, pinnae usually more than 2 cm wide, apex obtuse; scales at stipe bases dark brown in centre, gradually pale towards margin.
73a. Basal pinnules of pinnae often reflexed. Yunnan
71. A. lineare
73b. Pinnules patent or sloping upwards.
74a. Laminae 2-pinnate-pinnules pinnatisect to 3-pinnate-secondary pinnules pinnatilobate; basal pair of pinnae far bigger than the upper, oblong-ovate; costae and costules with short spine-like processes adaxially. Yunnan
72. A. chungtienense
74b. Laminae pinnate-pinnae pinnatisect to 2-pinnate- pinnules pinnatilobate; basal pair of pinnae similar to the upper ones or slightly shorter, narrowly lanceolate or narrowly oblanceolate; only in big pinnae costules with spine-like processes adaxially.
75a. The bases of middle pinnae asymmetrical; Teeth at pinnule or lobe margin sharp and long. Sichuan, Yunnan and Xizang
73. A. austro-orientale
75b. The bases of middle pinnae subsymmetric; Teeth at pinnule or lobe margin short. Sichuan, Guizhou, Yunnan and Xizang
74. A. dubium
44b. Indusia usually short-linear or oblong, straight, lateral on veinlets, often close to midrib, at most on laminae apex or acroscopic pinnule of pinna bases rarely curved; scales at stipe bases often black or dark-brown.
76a. Costae with subulate, short spine-like processes adaxially, costules or midribs without, occasionally with, but very short.
77a. Laminae broadly ovate or ovate, apex often cuspidate, occasionally long ovate, apex short acuminate; pinnae usually prominently stipitate (stipes usually more than 2--3 mm).
78a. Pinnules of basal 2--3 pairs of pinnae anadromic, pinnules prominently stipitate (stipes 3--5 mm); raches and costae glabrous abaxially. Anhui, Zhejiang, Taiwan, Hubei, Hunan, Fujian, Jiangxi, Guangxi, Sichuan and Guizhou
75. A. clivicola
78b. Only the pinnules of basal pair of pinnae anadromic, pinnules of rest pinnae subopposite or catadromic, shortly stipitate; raches and costae pubescent abaxially, rarely glabrate.
79a. Pinnules margin lobed, lobes may 1 mm, or duplicato-dentate. Guangxi
76. A. infrapuberulum
79b. Pinnules subentire or margin serrate or lobed.
80a. Pinnules lanceolate to narrowly lanceolate, length 3 or more times than width; sori subparallel to midrib; indusia erose. Shaanxi, Sichuan and Yunnan
77. A. criticum
80b. Pinnules deltoid, oblong-ovate or ovate, occasionally long ovate, length less than 3 times to width; sori not parallel to midrib; indusia entire or subentire.
81a. Laminae narrowly deltoid-ovate, narrow-ovate or ovate, pinnae 7 pairs or more except the pinnae of cuspidate apex; pinnules narrow, usually less than 5 mm wide at middle, subentire; pinna raches sparsely pubescent abaxially.
82a. Laminae ovato-oblong, cuspidate at apex; pinnae long (may up to 18 cm), pinnules more than 10 pairs. Hunan, Hubei, Shaanxi, Sichuan and Guizhou
78. A. brevistipes
82b. Laminae narrowly deltoid-ovate, acuminate at apex; pinnae short (5--9 mm), pinnules less than 10 pairs. Guangxi, Guangdong and Sichuan
79. A. kenzo-satakei
81b. Laminae deltoid or deltoid-ovate, pinnae about 5 pairs except pinnae of cuspidate apex; pinnules broad, usually more than 5 mm in width in the middle, serrate at margin; costae densely pubescent abaxially.
83a. The bases of basal pair of pinnae broadened, their basal pinnules biggest; pinnules auriculate at bases. Hunan and Guizhou
80. A. longius
83b. The basal pinnules of basal pair of pinnae shortened; pinnules auricled only at acroscopic bases.
84a. Pinnules of medial pinnae ascendant, oblong, serrulate at margin. Zhejiang, Anhui, Fujian, Jiangxi, Hunan, Hubei, Guangxi, Sichuan, Guizhou and Yunnan
81. A. wardii
84b. Pinnules of medial pinnae deltoid-broad-lanceolate, shallowly lobed to pinnatifid at margin . Hubei
82. A. venulosum
77b. Laminae oblong-ovate or lanceolate, occasionally ovate, acuminate at apex, occasionally cuspidate; pinnae usually sessile, or shortly stipitate (stipes less than 2 mm).
85a. Pinnules or lobes rounded-obtuse or truncate at apex, margin rounded-lobed or crenate; sori usually marginal. Guangxi and Yunnan
83. A. christensenii
85b. Pinnules or lobes acute or obtuse at apex, lobes or teeth at margin not crenate; sori usually close to costule.
86a. Basal pair of pinnules of pinnae overlay raches.
87a. Pinnae acuminate at apex; pinnule apex obtuse, margin teeth pointing directly outwards. Guangxi, Sichuan, Guizhou and Yunnan
84. A. delavayi
87b. Pinnae caudate-acuminate at apex; pinnule apex acute or obtuse, margin teeth sloping upwards.
88a. Pinnae long (14--16 cm), caudate-long acuminate at apex; pinnules short-acuminate at apex, long-auricled at acroscopic bases, midrib without spine-like processes adaxially. Sichuan and Hunan
85. A. caudiforme
88b. Pinnae short (10--12 cm), short-caudate at apex; pinnules acute at apex, short-auricled at acroscopic bases, midrib with spine-like processes adaxially. Taiwan
86. A. subrigescens
86b. Pinnae basal pair of pinnules not overlay raches.
89a. Scales at stipe bases ovato-lanceolate or lanceolate; pinnules of supra-medial pinnae anadromic or subopposite; sori short (about 1 mm), close to midrib, almost parallel to midrib. Yunnan
87. A. decorum
89b. Scales at stipe bases linear to linear-lanceolate; pinnules of supra- medial pinnae catadromic or subopposite; sori long (more than 1.5 mm), ascendant to midrib.
90a. Pinnae often shortly stipitate; pinnules narrowly oblong, broadly cuneate at bases; costae and raches with very short spine-like processes at apex adaxially. Yunnan
88. A. uniforme
90b. Pinnae sessile or very shortly stipitate; pinnules not narrowly oblong, cuneate at basiscopic bases, truncate and parallel to costae at acroscopic bases; spine-like processes of costae and raches at apex on the adaxial surface prominent.
91a. Laminae narrowly ovate or lanceolate, acuminate at apex.
92a. Costules prominently with spine-like processes adaxially; raches and costae densely pubescent or glabrous abaxially.
93a. Raches and costae glabrous abaxially; pinnules and lobes usually obtuse at apex, almost not lobed at margin, teeth thin. Yunnan
89. A. roseum
93b. Raches and costae densely pubescent abaxially; pinnules and lobes usually rounded-obtuse at apex, shallowly lobed or prominently serrate at margin. Yunnan, Sichuan and Taiwan
90. A. mengtzeense
92b. Costules without or with very short spine-like processes adaxially; raches and costae densely pubescent abaxially.
94a. Stipes, raches and costae pale purplish red; pinnae shortly stipitate (stipes 1--3 mm) or subsessile. Hubei, Hunan, Fujian, Guangdong, Guangxi, Sichuan, Guizhou, Yunnan and Taiwan
91. A. epirachis
94b. Stipes, raches and costae stramineous; pinnae sessile. Hubei, Hunan, Guangxi, Sichuan, Guizhou and Yunnan
92. A. pubicostatum
91b. Laminae ovate or deltoid-ovate, occasionally long ovate, slightly cuspidate or short acuminate at apex.
95a. Raches and costae glabrous abaxially; pinnae sessile.
96a. Basal pair of pinnae narrowly lanceolate, acroscopic pinnules of costae almost equal to the basiscopic ones. Hubei, Hunan, Anhui, Zhejiang, Fujian, Guangdong, Guangxi, Sichuan, Guizhou, Yunnan and Taiwan
93. A. otophorum
96b. Basal pair of pinnae ovato-lanceolate, basiscopic pinnules of costae longer than the acroscopic ones. Occur in Yunnan and Taiwan
94. A. kuratae
95b. Raches and costae pubescent abaxially; pinnae shortly stipitate.
97a. Spine-like processes of raches and costae short, sparsely glandular pubescent abaxially; pinnules separate, short-obtuse at apex. Taiwan
95. A. arisanense
97b. Spine-like processes of raches and costae long, densely with pale brown short glandular hairs abaxially; pinnules densely close, acute at apex. Yunnan
96. A. caudatum
76b. Costae and costules with long needlelike processes adaxially, ultimate lobe midrib often with too.
98a. Laminae 3-pinnate or more divided; secondary pinnules prominently stipitate; raches and costae glabrous abaxially.
99a. Laminae deltoid or ovato-deltoid; ultimate lobes broad, suboblong, usually more than 2--3 mm wide. Yunnan
97. A. adpressum
99b. Laminae elliptic; ultimate lobes slender and long, linear or long subulate, usually about 1 mm wide.
100a. Ultimate lobes linear, sloping upward patent; sori close to and along sides of costules, straight. Sichuan, Yunnan and Xizang
98. A. rhachidosorum
100b. Ultimate lobes long-subulate, pointing outwards patent; sori borne on ultimate lobes, often curved. Sichuan and Yunnan
99. A. araiostegioides
98b. Laminae pinnate to 2-pinnate; pinnules at most pinnatifid; raches and costae pubescent or glabrous abaxially.
101a. Sori oblong or short-linear, straight, close to midribs of pinnules or lobes, often partly overlap them when ripe; raches and costae glabrous abaxially, at most slightly pubescent at bases of costae.
102a. Small plants (fronds less than 25 cm), laminae narrowly deltoid or deltoid- ovate. (Three species under this item, the species 100. A. chingianum with long-creeping rhizome, laminae adaxial surface glabrous see above item 6) laminae adaxial surface glandular-pubescent.
103a. Laminae deltoid, basal pinnae not shortened; lobes subentire; sori close to costules and fall into two lines along sides of raches. Yunnan
101. A. supraspinenscens
103b. Laminae ovato-deltoid, basal pinnae a little shortened; lobe margin crenate or serrato-lobed; sori distant from costules. Sori distant from costules. Xizang
102. A. medogense
102b. Middle plants (fronds more than 30 cm), laminae broadly ovate to linear-lanceolate; laminae adaxial surface glabrous.
104a. Laminae pinnate; pinnae deltoid-lanceolate, less than 3.5 cm, at most pinnatisect, lobes densely neighboring. Yunnan
103. A. yui
104b. Laminae 2-pinnate; pinnae lanceolate or linear-lanceolate, 5--20 cm, pinnules separate from each other.
105a. Laminae ovate to oblong-lanceolate; ultimate lobe margin subentire or only slightly serrulate. Sichuan and Yunnan
104. A. fangii
105b. Laminae broadly ovate to ovate; ultimate lobe margin prominently with long or short teeth.
106a. Pinnae 13--16 pairs; pinnules oblong; raches and costae glabrous abaxially. Yunnan and Xizang
105. A. nyelamense
106b. Pinnae 6--7 pairs; pinnules obovate or oblong; raches and costae densely pubescent abaxially. Yunnan
106. A. yuanyangense
101b. Sori curved or straight, not close to midribs of pinnules or lobes, not overlap them when ripe; raches and costae pubescent abaxially, rarely glabrous.
107a. Laminae lanceolate to linear-lanceolate; pinnae usually more than 18 pairs; raches usually bearing bulblets below the apex, occasionally not.
108a. Laminae narrowly lanceolate to linear-lanceolate; many pairs of basal pinnae shortened; pinnae short (3--4 cm), subsessile, lobes less than 10 pairs, margin only with small teeth; bulblets of raches usually bigger. Guizhou and Yunnan
107. A. clarkei
108b. Laminae lanceolate to narrowly lanceolate; only 1--2 pairs of basal pinnae a little shortened; pinnae long (6--11 cm), usually shortly stipitate, lobes more than 10 pairs, biserrate or lobed at margin; bulblets of raches small.
109a. Pinnules deltoid-broadly lanceolate; lobes only with minute teeth at margin. Hunan, Jiangxi, Guangdong, Guangxi, Sichuan, Guizhou and Yunnan
108. A. viviparum
109b. Pinnules ovato-oblong; lobes with about 1 mm long teeth at margin. Hunan, Jiangxi, Guangdong, Guangxi, Sichuan, Guizhou, Yunnan, Xizang and Taiwan
109. A. strigillosum
107b. Laminae broadly ovate, ovate or lanceolate; pinnae less than 15 pairs; raches usually bearing no bulblets below apex, occasionally bearing.
110a. Small plants, less than 30 cm high; laminae lanceolate, less than 6 cm wide; pinnae narrow and small, less than 1 cm wide; sori borne on the base of veinlets. Yunnan, Xizang and Taiwan
110. A. nigripes
110b. Middle plants, usually more than 30 cm high; laminae broadly ovate or ovate, more than 6 cm wide; pinnae more than 1 cm wide; sori borne on the middle or infra-middle of veinlets.
111a. Pinnules pinnatilobate to pinnatifid, rarely pinnatipartite, apex obtuse, margin crenate; sori straight; laminae papyraceous when dry.
112a. Laminae oblong-ovate; basal several pairs of pinnae opposite or subopposite; stipes and raches stramineous. Yunnan and Guizhou
111. A. guangnanense
112b. Laminae subdeltoid; except basal pair subopposite, pinnae alternate; stipes and raches usually pale purplish red. Guangxi, Sichuan, Yunnan and Taiwan
112. A. silvicola
111b. Pinnules pinnatifid to pinnatisect, apex acute or obtuse, margin with sharp teeth; sori curved and straight or straight; laminae herbaceous when dry.
113a. Laminae ovate to narrowly ovate, occasionally lanceolate; pinnules or lobes separate from each other; pinnules oblong-ovate, two sides of midrib asymmetrical; sori straight, curved or other various in shape; raches often bearing bulblets below apex.
114a. Pinnules of medial pinnae anadromic. Taiwan, Zhejiang, Hunan, Hubei, Guangxi and Guizhou
113. A. cryptogrammoides
114b. Pinnules of medial pinnae subopposite. Jiangsu, Zhejiang, Anhui, Fujian, Taiwan, Jiangxi, Hunan, Hubei, Guangxi, Guangdong, Sichuan, Guizhou, Yunnan and Xizang
114. A. iseanum
113b. Laminae broadly ovate or ovate; pinnules or lobes close each other; pinnules oblong-ovate to long deltoid-ovate, two sides of midrib subsymmetric; sori straight, rarely curved; raches bearing no bulblets below apex.
115a. Stipes and raches pale purplish red, fragile; costae densely pubescent abaxially. Sichuan and Guizhou
115. A. imbricatum
115b. Stipes and raches stramineous, tough; costae glabrous abaxially.
116a. Stipes usually longer than laminae; laminae ovate, apex cuspidate; pinnae less than 10 pairs. Yunnan
116. A. xichouense
116b. Stipes usually shorter than laminae; laminae oblong, apex acuminate; pinnae about 13 pairs. Hunan
117. A. nanyueense
1. Athyrium wallichianum Ching in Bull. Fan Mem. Inst. Biol. Bot. ser. 8:497. 1938.
黑秆蹄盖蕨 hei gan ti gai jue
Aspidium brunonianum Wall. ex Mett.; Dryopteris brunoniana O. Ktze.; Lastrea brunoniana Presl; Nephrodium brunonianum Hook.
Rhizomes short, ascending, apices and stipes densely scaly; there are two kind of scales, the broad ones ovato-lanceolate, the narrow ones fibriform-tipped, both brown or somewhat chestnut-black, lustrous. Fronds caespitose; fertile fronds (14--)40--60 cm, stipes short, usually (4--)510(--15) cm long, (1--)3--4 mm thick, stipes and raches blackish-brown; lamina oblanceolate, (10--)20--50 cm long, (3--)5--7 cm broad at middle, obtuse at apex, narrowed toward base, bases 2--2.5 cm broad, pinnate, pinnae pinnatilobate to pinnatipartite; pinnae (12--)20--30 pairs, the lowers opposite, upward alternate, densely close, slightly ascendant, sessile, (2--)6--10 pairs of lower pinnae gradually shortened, the basal pinnae as ovato-triangular, only ca. 1 cm long, the middle pinnae oblong, 2--3.5 cm long, 1--1.5 cm broad, obtuse-rounded and shortly serrate at apex, symmetrical, near truncate, close to raches at base, pinnatipartite almost to costae; segments 6--8 pairs, patent, densely close, ovate, the basal pair a little bigger, 4--5 mm long and broad, duplicato-dentate, tooth triangular, acute at apex, patent, often revolute when dried. Veins visible abaxially, pinnate in segments, lateral veins 3--4 pairs, mostly simple, extending into teeth. Laminae thick-papery when dried, brown, glabrous, densely bearing red-brown or chestnut-black lanceolate and linear scales only along raches. Sori round or broadly ovate, dorsal on the acroscopic veinlets (but often on both veinlets of the basal acroscopic forked veinlets), 2--4 pairs per segment, in the middle between costules and margin; exindusiate. Spore perispore surface with folds, obscurely granulate.
In rock crevice under forests or alpine shrub-meadow; 3500--4800 m. Sichuan, NW Yunnan, SE Xizang [Nepal, Sikkim, Bhutan, N India, N Myanmar, N Pakistan, Kashmir]. Type from Nepal.
2. Athyrium dissitifolium (Bak.) C. Chr. in Contr. U.S. Nat. Herb. 26:296. t. 18. 1931.
疏叶蹄盖蕨 shu ye ti gai jue
Polypodium dissitifolium Bak. in Kew Bull. 1895:54. 1895. (Type P! Isotype K!); Athyrium drepanopterum A. Br. var. brevicaudatum Christ.; Dryopteris apicidens C. Chr.; Dryopteris cristulata Rosenst.; Athyrium fasciculatum Hand.-Mazz.; Dryopteris dissitifolia C. Chr.; Dryopteris incrassata C. Chr.; Phegopteris incrassata Christ; Polypodium apicidens Bak.
2a. var. dissitifolium
疏叶蹄盖蕨 shu ye ti gai jue
Rhizomes, short-creeping or ascending, densely scaly at apices and stipe bases; scales brown, linear-lanceolate, fibriform-tipped at apex. Fronds caespitose; fertile fronds 20--65 cm; stipes (5--)20--30 cm long, up to 2.5 mm thick, stramineous, lamina broadly lanceolate, 15--40 cm long, 5--12 cm broad, long-acuminate at apex, not narrowed at base, pinnate-pinnatilobate to 2-pinnate; pinnae 12--25 pairs, alternate, subpatent, nearly sessile, lanceolate, 2.5--14 cm long, 0.7--4 cm broad, long-acuminate at apex, bases truncate, pinnatipartite to pinnatisect at margin; pinnules or segments ca. 12 pairs, alternate, densely close each other, oblong, 0.5--2 cm long, 2--5 mm broad, obtuse with teeth at apex, entire or serrate at margin. Veins unclear adaxially, visible only abaxially, pinnate in pinnules or segments, lateral veins ca. 7 pairs, upswept, simple. Laminae thick-papery when dried, greenish brown, glabrous on both surfaces, raches and costae on the abaxial surface stramineous, glabrous. Sori round or elliptic, supra-medial on the veinlets, a little more close to margin, 5--6 pairs per pinnule or segment; exindusiate. Perispore surface with prominent folds. 2n=ca.80.
Under forests or in shrub grove roadside; 600--2700 m. Common. Hunan, Guangxi, Sichuan, Guizhou, Yunnan [N India, Nepal, Bhutan, Myanmar, Vietnam, Thailand]. Type from Yunnan (Mentzi).
2b. var. funebre (Christ) Ching et Z. R. Wang in Fl. Reip. Pop. Sin. 3(2):124. 1999.
二回疏叶蹄盖蕨 er hui shu ye ti gai jue
Athyrium drepanopterum (Kze.) A. Br. var. funebre Christ in Lecomte, Not. Syst. 1:46. 1909.
Pinnules obtuse at apex.
* In mixed forest margin; ca. 2300 m. Yunnan (Xi Shan, Kunming).
2c. var. kulhaitense (Atkinson ex Clarke) Ching in Ching & Hsieh in Acta Bot. Bor.-Occ. Sin. 6(1):22. 1986.
库尔海蹄盖蕨 ku er hai ti gai jue
Asplenium oxyphyllum Wall. ex Hook. forma kulhaitense W. S. Atkinson ex Clarke in Trans. Linn. Soc., Lond., 2. Bot. 1:493. 1880; Polypodium kulhaitense Atkinson ex Clarke, nom. inval., in syn.
It differs from the typical variety in laminae 2-pinnate; pinnae petioled; pinnules broadly lanceolate, acute at apex, bases asymmetrically cuneate, shallowly lobed or serrate at margin.
Under forests; 1300--2100 m. SW Yunnan [Nepal, N India, Myanmar].Type from Yunnan (Kulhaite).
3. Athyrium exindusiatum Ching in Ching et Hsieh in Acta Bot. Bor.-Occ. Sin. 6(2):102. 1986. (Type PE!)
无盖蹄盖蕨 wu gai ti gai jue
Athyrium drepanopterum auct. non (Kunze) A.Br. ex Milde 1867.
Fertile fronds up to 1 m; stipes ca. 50 cm long, 3--4 mm thick, bases clothed with brownish, lanceolate big scales, upward stramineous, glabrate; laminae oblong, ca. 65 cm long, 20--30 cm broad, long-acuminate at apex, not narrowed at base, 2-pinnate-pinnatifid; pinnae 20--22 pairs, alternate, ascendant, shortly petiolulate, 1.4 mm long, the lowest pinnae not shortened and not narrowed toward their base, lanceolate, (15--)25--28 cm long, 5--8 cm broad, long-acuminate at apex, bases rotund-cuneate, pinnate-pinnatifid; pinnules ca. 25 pairs, alternate, ascendant, nearly falcate-lanceolate, 3.5--4 mm long, bases 8--10 mm broad, acuminate at apex, asymmetrical at base, the acroscopic side truncate, somewhat auricled, parallel to costae, basiscopic side cuneate, connate with costae, pinnatilobate to pinnatipartite at margin (the basals pinnatipartite cut up to 2/3); segments 8--9 pairs, oblong, but the basiscopic one smaller, often deltoid, with long teeth at apex; upper pinnae lanceolate, long-acuminate at apex, basal acroscopic side truncate, parallel to raches, the basiscopic side broadly cuneate, sessile, slightly upward curved, biserrate at margin. Veins unclear adaxially, visible only abaxially, pinnate in segments, lateral veins 3--4 pairs, upswept, simple. Laminae papery when dried, glabrous, dark green adaxially, pale green abaxially. Raches and costae stramineous, glabrous abaxially, on the adaxial surface with very short spines. Sori round, dorsal on veinlets, 1--3 pairs per segment; exindusiate. Perispore surface with prominent folds.
In grassland under forests; 1000--2300 m. Yunnan, Xizang [India, Nepal, Myanmar].Type from Yunnan (Tengchong).
4. Athyrium pectinatum (Wall. ex Mett.) Bedd., Ferns S. Ind. 1:t. 155. 1863.
篦齿蹄盖蕨 bi chi ti gai jue
Asplenium pectinatum Wall., Num. List. no. 231,231.1, 1828, nom. nud. (Syntype K!); ex Mett., Abhandl. Senck. Naturf. Ges. (Frankfurt) 3(1):241. 1860; Asplenium filix-femina (L.) Bernh. var. pectinatum Clarke; Athyrium filix-femina (L.) Roth var. pectinatum Bedd.; Athyrium filix-femina (L.) Roth. ssp. pectinatum Lφve et Lφve.
Rhizomes slender, long, creeping, ascending at apex, densely clothed with dark-brown, lanceolate scales. Fronds approximate; fertile fronds 30--55 cm; stipes 15--25 cm long, 1--2 mm thick, brown-stramineous, clothed the same scales as rhizomes at bases, upward glabrate; laminae oblong-lanceolate, 15--35 cm long, 8--25 cm broad at middle, caudate-acuminate at apex, slightly narrowed at base, 3-pinnate to 3-pinnate-pinnatifid; pinnae ca. 20 pairs, the lowers opposite, upward subopposite, ascendant, shortly petiolulate(ca. 1.5 mm) or sub-sessile, the basal pair a little shortened, the middles deltoid-lanceolate, 10--15 cm, the lowers a little shortened, the middles deltoid-lanceolate, 10--15 cm long, ca. 3.5 cm broad et bases, acuminate at apex, truncate, the acroscopic side alongside raches at base, 2-pinnate; pinnules ca. 20 pairs, the basals opposite, upward alternate, patent, separate from each other, broadly lanceolate, up to 2.5 cm long, 8--10 mm broad at base, acute or acuminate at apex, symmetrical, rounded-truncate at base, pinnate; ultimate pinnules 8--9 pairs, the lowers opposite, upward alternate, sub-patent or ascendant, linear, 4--6 mm long, ca. 2 mm broad, obtuse at apex, decurrent and connate with the wings of costules at base, cut at margin, segments 3--4, teeth-like, upswept. Veins unclear adaxially, visible abaxially, pinnate in ultimate pinnules, lateral veins 3--4 pairs, upswept, veinlets simple, only 1 veinlet per tooth. Laminae near membranous when dried, yellow-green, and glabrous; raches, costae, costules (or midribs) stramineous, glabrous, with needle-shaped spines in the basiscopic margin of adaxial groove at costa or costule bases. Sori small, elliptic, 3--4 pairs per ultimate pinnule, 1 sorus per segment, alongside costules; indusia oblong, brown, membranous, erose at margin, persistent. Perispore surface with prominent folds. 2n=80.
Under coniferous forests; ca. 2100 m. SE Xizang [N India, Nepal, Sikkim]. Type from Nepal.
5. Athyrium ruilicola [ruilicolum] W. M. Chu in Acta Bot. Yunnan. Suppl. 5:40,41, t. 15. 1992. (Holotype, isotype PYU!).
瑞丽蹄盖蕨 rui li ti gai jue
Rhizomes creeping, densely clothed with brown, lanceolate scales. Fronds approximate; fertile fronds 23--40 cm; stipes 16--26 cm long, 0.5--1.5 mm thick, bases brown, sparsely bearing dark-brown lanceolate scales, upward stramineous, glabrate; laminae deltoid-ovate, 8--22 cm long, 3.5--8 cm broad, 2-pinnate; free pinnae 3--6 pairs, the basal pair not shortened or a little longer, subopposite, shortly petiolulate, sub-patent or slightly ascendant, ovate, 1.5--5.5 cm, the middle pinnae ovate-oblong, 1.5--4.5 cm long, 1--2 cm broad, acuminate or acute at apex, roundish at base, auricled at acroscopic side, very shortly petiolulate, alternate, upswept, pinnate; pinnules or segments 3--5 pairs, anadromic, deltoid-oblong, 0.1--1 cm long, 2--7 mm broad, obtuse at apex, shallowly toothed at margin, free pinnule nearly symmetrical at base, the segments on acroscopic side often auricled and parallel to costae, the basiscopic side broadly cuneate, slightly decurrent and connate to wings of costae. Veins visible on both surfaces, pinnate in pinnules, lateral veins 3--6 pairs, veinlets simple or forked. Laminae thin-herbaceous when dried, yellow-green, glabrous; raches and costae stramineous, glabrous abaxially. Sori oblong, J-shaped or horseshoe-shaped, across the basal acroscopic veinlets, biserial along two sides of costule; indusia bigger, same with sorus in shape, pallid, membranous, ciliated at margin, persistent. Spore perispore surface with folds.
* 1300--1400 m. Yunnan. Type from Yunnan (Ruili).
This species is similar to A. biserrulatum, but far thinner, fronds subdimorphic, stipes of sterile fronds shorter than lamina, stipes of fertile fronds far longer than lamina, lateral pinnae only 3--5 pairs, basal pair of pinnae not shortened or a little bigger.
6. Athyrium biserrulatum Christ in Bull. Acad. Int. Geogr. Bot. Mans 17:135--136. 1907 (Type P!).
苍山蹄盖蕨 cang shan ti gai jue
Asplenium filix-femina (L.) Bernh. var. polyspora Clarke; Athyrium filix-femina (L.) Roth. var. polysporum (Clarke) Bedd.; Athyrium polysporum (Clarke) Ching ex Mehra et Bir; Athyrium schimperi auct. non Moug. ex Fee, 1852.
Rhizomes slender, long, creeping, densely clothed with brownish, lanceolate scales. Fronds distant; fertile fronds 50--65 cm; stipes 15--20 cm long, 1.5--2 mm thick, bases brown, sparsely bearing dark-brown lanceolate scales, upward stramineous, glabrate; laminae oblong, 30--45 cm long, 14--18 cm broad, 2-pinnate; pinnae 15--20 pairs, basal 2--3(--4) pairs gradually shortened towards the base, opposite or subopposite, shortly petiolulate, sub-patent or slightly drooping, the basal pair triangular, ca. 2.5 cm, the middle pinnae lanceolate, 8--10 cm long, ca. 3 cm broad, acuminate at apex, bases roundish, shortly petiolulate, alternate, upswept, pinnate; pinnules 10--12 pairs, deltoid-oblong, 1--1.5 cm long, 5--7 mm broad, rounded and toothed at apex, somewhat asymmetrical at base, the acroscopic side often auricled and parallel to costa, the basiscopic side broadly cuneate, slightly decurrent, basal pinnules separate from each other widely, the uppers close each other, connate with wings of costae at base, shallowly lobed at margin, segments with spine-like teeth at apex. Veins visible adaxially, unclear abaxially, pinnate in pinnules, lateral veins ca. 6 pairs, veinlets simple or forked. Laminae thin-herbaceous when dried, pale green or dark green, glabrous; raches and costae on the abaxial surface stramineous, bearing pallid short hairs. Sori oblong, J-shaped or horseshoe-shaped, lateral on cross over basal acroscopic veinlets, biserial along two sides of costule (but often 2--4 on the basal acroscopic segment); indusia bigger, same with sorus in shape, pallid, membranous, ciliated at margin, persistent. Perispore with folds, which connected into large reticula. 2n=80.
Under forests or forest margin; 2000--3000 m. Sichuan, W Guizhou, C-NW Yunnan, SE Xizang [N India, Pakistan, Sikkim, Bhutan, Nepal, Myanmar]. Type from Yunnan (Cang Shan, Dali).
This species is very similar to African Athyrium schimperi Moug. ex Fee. The type of the later was collected from Ethiopia. Might be they are subspecies.
7. Athyrium wumonshanicum Ching in Ching et Hsieh in Acta Bot. Bor.-Occ. Sin. 6(1):20. 1986. (Type PE! Isotype PYU!).
乌蒙山蹄盖蕨 wu meng shan ti gai jue
Rhizomes slender, long, creeping, clothed with brown, narrowly lanceolate scales at apices and stipe bases. Fronds approximate; fertile fronds up to 60 cm; stipes 17 cm long, ca. 2 mm thick at base, brown, upward stramineous, glabrate; laminae ovato-oblong, 40 cm long, ca. 16 cm broad at middle, acuminate at apex, almost not narrowed at base, 3-pinnate; pinnae ca. 14 pairs, opposite, ascendant, shortly petiolulate(ca. 2.5 mm), the basal pair almost not shortened, the second pair of pinnae oblong-lanceolate, up to 13 cm long, ca. 5 cm broad, acuminate at apex, bases sub-truncate, more or less alongside raches, 2-pinnate; pinnules ca. 12 pairs, alternate, sub-patent, sessile, oblong-lanceolate, up to 2.5 cm long, bases ca. 1 cm broad, acuminate at apex, rotund-cuneate, slightly decurrent and more or less connate with wings of costae, pinnatipartite, almost cut to costules at margin; ultimate pinnules 5--6 pairs, ascendant, oblong-lanceolate, the basals up to 7 mm long, ca. 3 mm broad, upward gradually shortened, acute and with 3--4 long spine-like teeth at apex, toothed at margin. Veins thick, slightly depressed adaxially, protuberant abaxially, pinnate at ultimate pinnules, lateral veins 4 pairs, upswept, simple. Laminae herbaceous when dried, pale-green, sparsely bearing pale hairs adaxially, on the abaxial surface of costae and costules sparsely bearing short hairs. Sori big, oblong, 26 sori per ultimate pinnule; indusia bigger, same with sorus in shape or J-shaped, pale-brown, membranous, unequally toothed at margin, persistent. Perispore surface prominently with folds.
* About 1800 m. N Yunnan (Wumeng Shan, Luquan).
This species is similar to Athyrium biserrulatum in shape. However, its lamina 3-pinnate, bearing short hairs adaxially, with aristate teeth at margin.
8. Athyrium niponicum (Mett.) Hance in Journ. Linn. Soc. 13:92. 1873.
日本蹄盖蕨 ri ben ti gai jue
Asplenium niponicum Mett. in Ann. Mus. Bot. Lugd. Bat. 2:240. 1866; Asplenium uropteron Miq.; Athyrium uropteron C. Chr.; Asplenium niponicum var. α minus, β uropteron etγ longipes Franch. et Sav.; Athyrium biondii Christ; Athyrium faberi Christ; Athyrium oshimense Christ; Athyrium yunnanense Christ; Athyrium fissum Christ; Athyrium sylvestrii Christ; Athyrium matsumurae Christ ex Matsum.; Athyrium pachyphlebium C. Chr.; Athyrium niponicum (Mett.) Hance var. pachyphlebium (C. Chr.) Kitag.
Form. 1 Athyrium niponicum form. niponicum
日本蹄盖蕨 ri ben ti gai jue
Summer-green. Rhizomes short-creeping, ascending, densely clothed with brownish, narrowly lanceolate scales at apex and stipe bases. Fronds caespitose; fertile fronds (25--)30--75(--120) cm; stipes 10--35(--50) cm long, (1.5--)2--3(--5) mm thick at base, blackish brown, upward stramineous, sparsely bearing small scales; scales lanceolate, entire at margin, pale brown; laminae ovato-oblong, (15--)23--30(--70) cm long, (11--)15--25(--50) cm broad at middle, abruptly narrowed at apex, broadly rounded at base, 2-pinnate to 3-pinnate; pinnae under abruptly narrowed apex 5--7(--14) pairs, alternate, ascendant, petiolate(3--15 mm), slightly curved upwards, the basal pair a little longer and bigger, oblong-lanceolate, (5--)7--15(--25) cm long, (2--)2.5--6(--12) cm broad at middle, abruptly narrowed, long-acuminate, somewhat caudate at apex, broadly cuneate or rounded at base, middle pinnae lanceolate, pinnate to 2-pinnate; pinnules (8--)12--15 pairs, alternate, ascendant or patent, shortly petiolulate or sub-sessile, usually broadly lanceolate or oblong-lanceolate, also lanceolate, the middles 1--4(--6) cm long, bases 1--2 cm broad, acuminate at apex, asymmetrical at base, the acroscopic side sub-truncate, auricled, parallel to costae, the basiscopic side cuneate, dentate or pinnatipartite almost cut to the wings of costules at margin; segments 8--10 pairs, lanceolate, oblong or linear-lanceolate, acute at apex, deeply serrate at margin. Veins visible abaxially, pinnate in segments, lateral veins 4--5 pairs, upswept, simple. Laminae herbaceous or thin-papery when dried, pale-green or yellow-green, glabrous on both surfaces; raches and costae on the abaxial surface pale red-purplish, sparsely bearing brownish linear small scales. Sori oblong, hooked, J-shaped or horseshoe-shaped, 4--12 pairs per ultimate segment; indusia same with sori in shape, brown, membranous, somewhat erose at margin, persistent or partly caducous. Perispore surface with prominent low folds. 2n=80.
Under forests, streamside, shaded and wet mountain slope, shrub or grassy slopes; Near sea level to 2600 m. Liaoning, Hebei, Shanxi, Shaanxi, Ningxia, Gansu, Shandong, Jiangsu, Anhui, Taiwan, Zhejiang, Jiangxi, Henan, Hubei, Hunan, Guangdong, Guangxi, Sichuan, Chongqing, Guizhou, Yunnan [Japan, Korea, Nepal, Vietnam, Myanmar]. Type from Japan.
Form. 2. Athyrium niponicum form. cristato-flabellatum (Makino) Nemegata et Kurata, Enum. Jap. Pterid. 279.338. 1961.
鸡冠蹄盖蕨 ji guan ti gai jue
Athyrium niponicum var. cristato-flabellatum Makino; Athyrium niponicum monstr. cristato-flabellatum Nakaike.
Pinnae or segments narrowed as flattened cristate at apex.
Cultivated; Shandong (Qingdao).
9. Athyrium brevisorum (Wall. ex Hook.) Moore, Ind. Fil. 117. 1859.
中缅蹄盖蕨 zhong mian ti gai jue
Asplenium brevisorum Wall., nom. nud. 1828, ex Hook. Sp. Fil. 3:229. 1860.
Rhizomes short-creeping, ascending. Fronds caespitose; fertile fronds 80--90 cm; stipes 45--55 cm long, 2--3 mm thick, bases brown, sparsely bearing brown, lanceolate or linear-lanceolate scales, upward stramineous, glabrate; laminae ovate or oblong-ovate, 35--45 cm, up to 25 cm broad at middle, abruptly narrowed at apex, not narrowed at base, 2-pinnate; pinnae ca. 12 pairs, alternate, ascendant, petioled (4--8 mm), the basal pair not shortened, lanceolate, ca. 19 cm long, 3--4 cm broad, long-acuminate at apex, rounded-truncate at base, or the acroscopic side parallel to raches, the basiscopic side narrowly cuneate, pinnate; pinnules 16--18 pairs, upswept, lanceolate, 2--3 cm long, 8 mm broad at middle, acuminate at apex, connate with costae at base, pinnules dentate at margin (sometimes basal pinnules shallowly lobed, segments with 2--3 small teeth at apex); abruptly narrowed part of lamina at apex very long pinna-like, pinnatipartite almost up to costae. Veins visible on both surfaces, pinnate in pinnules, lateral veins forked. Laminae herbaceous when dried, pale yellow-brown, raches and costae sparsely bearing pallid unicellular glandular-hairs abaxially. Sori short-linear or J-shaped, lateral on or cross over acroscopic veinlets, 8--10 pairs per pinnule (if pinnules pinnatilobate, 2--3 pairs per segment); indusia bigger, same with sorus in shape, pale-brown, membranous, slightly erose at margin, persistent. perispore surface with prominent folds.
Under forests; 800--1500 m. W Yunnan (Valley of Yingjiang) [Myanmar, Pakistan]. Type from Nepal.
10. Athyrium yokoscense (Franch. et Sav.) Christ in Bull. Herb. Boiss. 4:668. 1896.
禾秆蹄盖蕨 he gan ti gai jue
Asplenium yokoscense Franch. et Sav., Enum. Pl. Jap. 2:225. 1877. (Syntype, lectotype P!); Athyrium coreanum Christ(Syntype, pro parte P!); Athyrium demissum Christ; Athyrium flaccidum Christ; Athyrium pachysorum Christ; Aspidium subspinulosum Christ; Dryopteris subspinulosa C. Chr.
Rhizomes short, stout, erect, densely clothed with yellow-brown, narrowly lanceolate scales at apex. Fronds caespitose; fertile fronds (30--)40--60 cm; stipes (10--)12--20(--25) cm long, ca. 2.5 mm thick, bases dark brown, densely clothed the same scales as rhizomes, upward stramineous, glabrate; laminae oblong-lanceolate, 18--45 cm long, (8--) 11--15 cm broad, acuminate at apex, not narrowed at base, pinnate-pinnatipartite to 2-pinnate, pinnules shallowly lobed; pinnae 12--18 pairs, the lowers subopposite, upward alternate, patent or slightly ascendant, sessile, lanceolate, the middles (3.5--)7--9 cm long, (1.2--)1.5--2 cm broad, long-acuminate at apex, the acroscopic side truncate, the basiscopic side cuneate at base, pinnate; pinnules ca. 12 pairs, oblong-lanceolate, up to 1 cm long, ca. 5 mm broad, acute at apex, auricled at acroscopic base, decurrent and often connate to wings of costae at basiscopic base, shallowly lobed or dentate at margin, segments with 2--3 short teeth at apex. Veins visible abaxially, pinnate in pinnules, lateral veins forked. Raches and costae on the abaxial surface stramineous, slightly bearing brownish, lanceolate small scales, short thick spine-like processes present in the basiscopic margin of adaxial groove at costa or costule bases. Sori rotund or elliptic, medial on veinlets; indusia elliptic, J-shaped or horseshoe-shaped, brownish, membranous, entire, persistent. Perispore surface with prominent folds. 2n=80.
In rock crevice under forests; 100--2400 m. Heilongjiang, Jilin, Liaoning, Shandong, Jiangsu, Anhui, Zhejiang, Jiangxi, Henan, Hunan, Chongqing, NE Guizhou [Korea, Japan, E Russia].Type from Japan (Yokosuka).
According to the report in Key to the Plants from NE China (Ed. By Peiyun Fu), there is a variety in China: Athyrium yokoscense (Franch. et Sav.) Christ var. kirisimaense (Tagawa) Li et J. Z. Wang in Peiyun Fu (ed.), Clavis Plantarum Chinae Boreali-Orientalis, Editio Seunda. 36, Beijing: Science Press, 1995. (─ Athyrium kirisimaense Tagawa in Acta Phytotax. Geobot. 2:22. 1933.) It differs from the typical variety in laminae pinnate to 2-pinnate-pinnatifid, most pinnules petioled, scales in rhizomes lanceolate to ovate-lanceolate. Type from Japan. If it is a good variety and occurs in China, it is still waiting for further investigation.
11. Athyrium fauriei (Christ) Makino in Bot. Mag. Tokyo 17:160. 1903.
佛瑞蹄盖蕨 fo rui ti gai jue
Nephrodium fauriei Christ in Bull. Herb. Boiss. 4:671. 1896. (Type P! Isotype K!); Aspidium fauriei Christ; Athyrium yokoscense Christ var. fauriei Tagawa.
This species is similar to Athyrium yokoscense, but its lamina pinnate, pinnae far broader and larger, pinnatilobate to pinnatipartite, scales red-brown, blackish-brown in centre, raches subglabrous, pinnae sparsely with short hairs only at bases, indusia large.
China [Japan, Philippines]. Type from Japan.
12. Athyrium fallaciosum Milde, Fil. Europe. & Atlant. 6:54. 1867.
麦秆蹄盖蕨 mai gan ti gai jue
Asplenium mongolicum Franch.; Athyrium mongolicum Diels var. purdonii C. Chr.
Rhizomes short-creeping, ascending at apex, densely clothed with dark-brown, subulate lanceolate scales. Fronds caespitose; fertile fronds 30--50 cm; stipes 5--7 cm long, 1--2 mm thick, a little swollen, brown, densely clothed with the same scales at base, gradually becoming glabrate upwards, stramineous; laminae near oblanceolate, 25--40 cm long, 6--8 cm broad, acuminate at apex, narrowed toward base, pinnate, pinnae pinnatipartite; pinnae (segments) 20--24 pairs, 6--7 pairs of the lowers gradually shortened as triangular， auriculate small lobes, subopposite, sessile, patent or slightly inclined, the basal pair 1--1.2 cm long, ca. 7 mm broad; middle pinnae broadly lanceolate, 3--4 cm long, 1.2--1.5 cm broad at base, obtuse or acute at apex, truncate at base, sessile, patent, slightly curved up, pinnatipartite up to costae; segments ca. 8 pairs, the basal acroscopic one larger, 7--8 mm long, ca. 5 mm broad, oblong, obtuse-rounded and crenate at apex, dentate at margin, upward segments shorter, densely close. Veins visible on both surfaces, pinnate in segments, lateral veins 3--4 pairs, simple, occasionally forked. Laminae herbaceous when dried, green or greenish brown, glabrous; raches stramineous, occasionally bearing brown lanceolate scales. Sori big, mostly round-reniform or horseshoe-shaped, 2--3 pairs per segment; indusia bigger, same with sorus in shape, membranous, pallid, ciliated or erose at margin, persistent. Spore perispore with low folds. 2n=80@.
In valley, under forests, or wet rock crevice; 1200--2200 m. Heilongjiang, Jilin, Liaoning, Nei Mongol, Hebei, Shanxi, Shaanxi, Ningxia, Gansu, Henan, Hubei, N Sichuan [Korea, Japan]. Type from Beijing.
13. Athyrium rupicola (Edgew ex Hope) C. Chr., Ind. Fil. 1:145. 1906.
岩生蹄盖蕨 yan sheng ti gai jue
Asplenium rupicola Edgew ex Hope in Journ. Bomb. Nat. Hist. Soc. 12:531--532. t. 5. 1899. (Syntype P! BM!); Asplenium filix-femina (L.) Bernh. var. retusa subvar. elongata Clarke ex Blanf.
Rhizomes short-creeping, ascending at apex, densely clothed with chestnut-coloured or dark-brown, long subulate lanceolate, fibriform-tipped scales. Fronds caespitose; fertile fronds 25--55 cm; stipes 5--15 cm long, 1.2--2.5 mm thick, bases brown, densely clothed the same scale as rhizomes, upward stramineous, glabrate; laminae narrowly or broadly lanceolate, 20--35 cm long, 5--8.5 cm broad, acuminate at apex, gradually narrowed toward base, pinnate-pinnatipartite; pinnae 20--30 pairs, many pairs of the lowers gradually shortened as triangular-auricled, ca. 1 cm long, subopposite, slightly inclined, the middles narrowly deltoid-lanceolate or broadly lanceolate, 2.5--4 cm long, 8--16 mm broad, acuminate at apex, bases truncate, nearly opposite or alternate, sessile, patent or slightly curved upward, pinnatipartite to the narrow wings of costae; pinnules 8--12 pairs, the basal acroscopic one larger, the rest oblong, 4--5 mm long, 1.5--3 mm broad, obtuse or acute at apex, serrate or biserrate at apex, connate with costae and joining each other by narrow wing at base, serrate or biserrate at margin. Veins unclear adaxially, visible abaxially, pinnate in segments, lateral veins ca. 6 pairs, veinlets simple or forked. Laminae herbaceous when dried, greenish brown, glabrous on both surfaces; raches and costae on the abaxial surface stramineous, glabrous, or occasionally bearing a few brownish, fibrous scales. Sori round, dorsal on acroscopic veinlets of segments, 2--5 pairs per pinnule; indusia big, round-reniform or near ovate, pale-brown, erose at margin, persistent. Perispore with prominent folds, surface granuliferous. 2n=80.
In valley under forests, rock crevice or roadside damp area; 1800--3800 m. Sichuan, Yunnan, SE Xizang [Nepal, Sikkim, Bhutan, Kashmir, India, N Pakistan]. Type from NW India.
14. Athyrium glandulosum Ching in Ching et Hsieh in Acta Bot. Bor.-Occ. Sin. 6(1):19--20. 1986 (Type PE!).
腺毛蹄盖蕨 xian mao ti gai jue
Rhizomes short, erect or ascending, densely clothed with red-brown, narrowly lanceolate scales at apices and stipe bases. Fronds caespitose; fertile fronds ca. 50 cm; stipes 14--16 cm long, ca. 2 mm thick at base, dull brown, upward brownish stramineous, glabrate; laminae lanceolate, 30--35 cm long, ca. 6 cm broad at middle, acuminate at apex, gradually narrowed toward base, pinnate-pinnatifid; pinnae ca. 25 pairs, alternate, ascendant, sessile, 4--5 pairs of the lowers gradually shortened, inclined, the basal pair ca. 1 cm long, middle pinnae falcate-lanceolate, 3--4 cm long, 1.2--1.8 cm broad at base, acuminate at apex, sub-truncate at base, the acroscopic side slightly auricled, pinnatipartite almost cut to costae; segments ca. 10 pairs, slightly ascendant, close, oblong-lanceolate, 7--8 mm long, ca. 2 mm broad, acute at apex, decurrent to narrow costal wing at base, 4--5 pairs lobed or dentate at margin. Veins visible on both surfaces, pinnate in pinnules, lateral veins 4--5 pairs, very upswept, simple or forked. Laminae herbaceous when dried, brown, glabrous on both surfaces; raches and costae brown-stramineous abaxially, densely bearing pale, short glandular-hairs. Sori round, medial on veinlets, 4--5 pairs per pinnule; indusia large, broadly ovate, pale, membranous, sub-erose at margin, persistent. Perispore surface with prominent folds.
* Forest margin or shrub groves; 1500--3500 m. Sichuan, NW Yunnan. Type from Yunnan (Heqing).
This species is quite similar to Athyrium rupicola in shape, but raches and costae densely bearing pale, short glandular-hairs abaxially, middle pinnae falcate-lanceolate. So it can be easily recognized.
15. Athyrium minimum Ching in Ching et Hsieh in Acta Bot. Bor.-Occ. Sin. 6(3):151. 1986. (Type PE!)
小蹄盖蕨 xiao ti gai jue
Rhizomes short small, erect, sparsely bearing brownish, lanceolate scales at apices and stipe bases. Fronds caespitose; fertile fronds ca. 10 cm; stipes 2--3 cm long, ca. 0.5 mm thick, stramineous; laminae lanceolate, 4--6 cm long, 1--1.5 cm broad, acuminate at apex, almost not narrowed at base, pinnate-pinnatifid, petioled (ca. 1 mm), patent or ascendant, the basal pair a little shortened, near ovate or ovato-triangular, reflexed; middle pinnae unequally ovate or rhomboid, 8--10 mm long, 3.5--4 mm broad, obtuse at apex, asymmetrical at base, the acroscopic side auricled (auricle free, broadly ovate or oblong, with triangular teeth at margin), broadly cuneate at basiscopic base, pinnatipartite almost cut to costae at margin; segments small, 2 pairs, obovate or oblong, with 12 sharp teeth at apex. Veins visible abaxially, pinnate in segments, lateral veins forked, 1 veinlet per tooth. Laminae papery when dried, glabrous, dark brown adaxially, pale brown abaxially; raches and costae stramineous abaxially, occasionally with 1 or 2 brownish, linear small scales. Sori J-shaped or horseshoe-shaped, across on acroscopic veinlets, only 1 sorus per segment, but sometimes 2 on the auriculate lobes; indusia same with sori in shape, brownish, membranous, erose at margin, persistent. Perispore surface without folds.
* In valley under forests, wet places. Taiwan (Danshui).
16. Athyrium nakanoi Makino in Bot. Mag. Tokyo 23:247. 1909.
红苞蹄盖蕨 hong bao ti gai jue
Asplenium macrocarpum var. unipinnatum Clarke; Athyrium macrocarpum var. unipinnatum Bedd.; Nephrolepis tenuissima Hay.; Athyrium tenuissimum Kodama; Athyrium obtusifolium Rosenst.
Evergreen. Rhizomes erect or short-ascending, clothed with dark-brown, lanceolate scales at apices and stipe bases. Fronds caespitose; fertile fronds (10--)30--38(--50) cm; stipes 8--14(--18) cm long, 1--1.5 mm thick at base, dark-brown, upward stramineous, clothed with blackish brown short glandular-hairs and brown small scales; laminae lanceolate, (7--)22--28(--32) cm long, (2.5--)3--4.5(--7) cm broad, acuminate at apex, slightly narrowed at base, pinnate; pinnae 16--18 pairs, alternate, subpatent, petioled (ca. 1 mm), the basal 1--2 pairs a little shortened, inclined or recurved, middle pinnae oblong, (1--)2--2.5(--3.5) cm long, (5--)6--7(--12) mm broad, obtuse at apex, shortly toothed, extremely asymmetrical at base, the acroscopic side prominently auricled, the auricles lingulate-oblong or subtriangular, 4--7 mm, obtuse at apex, entire or somewhat repand to dentate, the basal basiscopic side of pinnae cuneate, deltoid-lobed or repand-teethed at margin. Veins unclear adaxially, visible abaxially, pinnate in pinnae, lateral veins 8--10 pairs, usually veinlets forked, but pinnate at auriculate lobes. Laminae papery when dried, yellowish brown or greenish brown, glabrous on both surfaces; raches stramineous, sparsely bearing blackish-brown or dark brown short glandular-hairs. Sori large, mostly horseshoe-shaped or round-reniform, in some cases J-shaped at apex, supramedial on the acroscopic veinlets, biserial along two sides of costule, frequently 25 sori at auriculate lobes; indusia bigger, same with sorus in shape, brown, membranous, erose at margin, persistent. Perispore with a few folds, slightly spinulate on the surface. 2n=160.
Under evergreen broadleaf forests and shrub groves, on rocks or streamside in valley; 1300--3400 m. Taiwan, Yunnan, SE Xizang [Japan, Bhutan, Nepal, N India]. Type from S Japan (Kyushu).
17. Athyrium zhenfengense Ching in Ching et Hsieh in Acta Bot. Bor.-Occ. Sin. 6(3):151. 1986. (Type PE!)
贞丰蹄盖蕨 zhen feng ti gai jue
Rhizomes short, stout, erect, clothed with brown, lanceolate scales at apices and stipe bases. Fronds caespitose; fertile fronds 20--25 cm; stipes 6--10 cm long, 1--1.5 mm thick at base, brown, upward brown-stramineous, sparsely bearing small scales and brownish short glandular-hairs; laminae lanceolate, 14--17 cm long, 3--4 cm broad at middle, acuminate at apex, slightly caudate, not narrowed at base, pinnate; pinnae 13--15 pairs, alternate, ascendant, stalked (1--1.5 mm), the basal 1--2 pairs similar to the upper neighbours (falcate-oblong), 2--2.5 cm long, 6--8 mm broad at middle, shortly acute and shortly toothed at apex, very asymmetrical at base, the acroscopic side auricled, parallels to raches, the auricles 6--7 mm long, ca. 3 mm broad, rounded and with a few small teeth at apex, the basiscopic side of pinnae cuneate, cut into triangular teeth at margin. Veins visible adaxially, clearly visible abaxially, pinnate or 2- to 3-forked on auricles or segments, veinlets simple. Laminae papery when dried, glabrous, brown adaxially, yellow-green abaxially; raches brown-stramineous, sparsely bearing brownish small scales and short glandular-hairs. Sori nearly round, inframedial on the acroscopic veinlets, 1--3 sori per segment, often 2--3 pairs on auriculate lobes; indusia J-shaped or horseshoe-shaped, dark-brown, membranous, erose at margin, persistent. Perispore surface with prominent folds.
* In valley under forests; 1500--1600 m. SW Guizhou (Zhenfeng).
This species is very similar to Athyrium nakanoi, but pinnae acute at apex, auriculate lobes with small teeth at apex, the short glandular-hairs on raches brown , but not blackish or dark brown.
18. Athyrium bucahwangense Ching in C. Y. Wu, Fl. Xizang. 1:132--133. 1983 (Type PE!).
圆果蹄盖蕨 yuan guo ti gai jue
Rhizomes short, stout, ascending, clothed with brown, lanceolate scales at apex. Fronds caespitose; fertile fronds 20--80 cm; stipes up to 25 cm long, 2.5--3 mm thick, brown, clothed with the same scales as rhizomes at bases, upward glabrate; lamina ovato-triangular, ca. 35 cm long, ca. 30 cm broad at base, short-acuminate at apex, bases broadly ovate, 2-pinnate to 3-pinnate-pinnatifid; pinnae 10--12 pairs, alternate, ascendant, petioled (5--12 mm), basal 2--3 pairs of pinnae similar each other in size, oblong-lanceolate, 15--16 cm long, 5--8 cm broad, long-acuminate at apex, bases symmetrical, truncate, 2-pinnate-pinnatifid; pinnules ca. 15 pairs, alternate, patent, petiolate(1--2 mm), deltoid-lanceolate, the middles 2.3--4 cm long, bases 1.4--2 cm broad, obtuse at apex, bases asymmetrical, the acroscopic side auricled, the basiscopic side cuneate, pinnate; secondary pinnules 7--8 pairs, the basal acroscopic one larger, ovate, 8--9 mm, pinnatilobate, the rest oblong, ca. 4 mm long, 1.5--2.5 mm broad, obtuse-rounded at apex, subentire or slightly toothed at margin; segments entire or sparsely crenulate with 12 teeth at margin; pinnae gradually narrower upward, their lower pinnules not shortened, the basal acroscopic pinnule the largest， parallel to or partly overlapping raches，divided almost same with the lower pinnae; the topic pinnae lanceolate, auricled at acroscopic base. Veins more or less visible adaxially, clearly visible abaxially, pinnate on ultimate pinnules, lateral veins simple or forked on the basal acroscopic side. Laminae herbaceous when dried, blackish brown adaxially, pale brown abaxially. Raches and costae brown, glabrous. Sori small, round, dorsal on the basal acroscopic veinlet top or lateral supramedial on the acroscopic veinlets, 1 sorus per segment; indusia round-reniform, elliptic or horseshoe-shaped, brown, membranous, entire, persistent. Perispore surface with a few folds.
* Under coniferous forests or shrub groves; 1900--3800 m. NW Yunnan, SE Xizang. Type from Yunnan (Gongshan, Chalong-Chaluo Watershed).
19. Athyrium erythropodum Hay. Ic. Pl. Form. 4:233, f. 163. 1914. (Type TI!).
红柄蹄盖蕨 hong bing ti gai jue
Rhizomes moderately stout, erect, clothed with pale-brown lanceolate scales at apex. Fronds caespitose; fertile fronds ca. 70 cm; stipes (8--)15--40 cm long, 1--2 mm thick, bases sparsely bearing lanceolate scales; laminae ovato-lanceolate, 25--35 cm long, 10--23 cm broad, somewhat abruptly narrowed and acuminate at apex, rounded at base, 2-pinnate, pinnules pinnatipartite; pinnae ca. 8 pairs, ovato-lanceolate, 10--15 cm long, 2.5--5 cm broad, petiolule 5--7 mm, the basal pair subopposite, upward alternate, upswept, the basal pair in the same size as the second pair, long-acuminate at apex, nearly rounded, symmetrical at base; pinnules ca. 12 pairs, sessile, separated each other by similar space, ascendant, ca. 3 cm long, 7 mm broad, deltoid-oblong, short-acuminate at apex, the acroscopic slightly elongate, parallel to costae, the basiscopic side cuneate at base, pinnatipartite almost cut to costules at margin; segments ca. 6 pairs, oblong, upswept, the basal acroscopic one a little bigger, 8 mm long, 3 mm broad, oblong, acute at apex. veins pinnate in segments, veinlets simple, sometimes forked at the basal big segments. Laminae herbaceous when dried, raches on the adaxial surface without spine-like processes. Sori oblong, in some cases hooked, sitting on acroscopic veinlets in segments, 1--3 sori per segment; indusia same with sori in shape, membranous, somewhat erose at margin, persistent.
* Taiwan [S Japan]. Type from Taiwan.
20. Athyrium drepanopterum (Kunze) A. Br. ex Milde in Milde, Fil. Europe. Atlant. 49. 1867.
多变蹄盖蕨 duo bian ti gai jue
Polypodium drepanopterum Kunze in Linnaea 23:278,318. 1850; Asplenium drepanopterum A. Br.; Aspidium drepanopterum A. Br. ex Mett.; Aspidium eburneum Wall. ex Kunze; Asplenium eburneum J. Sm.; Athyrium eburneum J. Sm.; Aspidium drepanopterum A. Br. var. decompositum Christ; Athyrium oxyphyllum Moore; Asplenium oxyphyllum Hook.; Athyrium niponicum Hance var. elatius Christ; Athyrium veitchii Christ; Athyrium woodsioides Christ; Athyrium moupinense Christ; Dryopteris takeoi Hay.; Athyrium micans Tagawa.
Rhizomes short, ascending, densely clothed with brown subulate lanceolate scales at apex. Fronds caespitose; fertile fronds (10--)25--90 cm; stipes (2.5--)10--30 cm long, 1.5--3 mm thick, bases blackish brown, densely clothed with the same scales as rhizomes, upward stramineous; laminae oblong-lanceolate, (10--)17--55 cm long, 5--30 cm broad at middle, acuminate at apex, pinnate to 2-pinnate, pinnules pinnatisect. Pinnae ca. 16 pairs, alternate, petioled (1--4 mm), basal pinnae not shortened, patent, middle pinnae oblong-lanceolate or broadly lanceolate, acuminate or long-acuminate at apex, asymmetrical at base, the acroscopic side auricled, parallel to raches, the basiscopic side rounded-truncate, pinnate to pinnate-pinnatipartite; pinnules 6--10 pairs, alternate, ascendant, close each other or separated each other by narrow space, the basal acroscopic one the biggest, deltoid-oblong, pinnatilobate or pinnatipartite, rest pinnules oblong or oblong-lanceolate, 7--15 mm long, bases 3--7 mm broad, acute and shortly teethed at apex, asymmetrical, the acroscopic side auricled, parallel to raches, connate with costae at base, subentire or pinnatisect almost cut to costae at margin; segments oblong, 5--6 mm long, 2--3 mm broad, acute at apex, with 1--2 small teeth at top. Veins unclear, lateral veins pinnate, veinlets simple. Laminae near herbaceous when dried, glabrous, dark green, lustrous adaxially. Sori near round or round-reniform, dorsal, medial on acroscopic veinlets, 1--5 pairs per segment; indusia small, round-reniform or horseshoe-shaped, brown, membranous, erose at margin, often partly fall off. Perispore surface prominently with low folds. 2n=80,160.
In valley, under forests, or in wet granite crevice; 700--2300 m. Taiwan, W Sichuan, Guizhou, C Yunnan, Xizang [N India, Nepal, Bhutan, Myanmar, Vietnam, Philippines].
This species is very similar to Athyrium dissitifolium var. kulhaitense, but laminae lustrous adaxially, with indusia, so it is easily to be distinguished.
21.Athyrium fimbriatum (Wall. ex Hook.) Moore, Ind. Fil. 185. 1860.
喜马拉雅蹄盖蕨 hi ma la ya ti gai jue
Asplenium fimbriatum Wall. ex Hook. Sp. Fil. 3:234. 1860, non Kunze 1844; Aspidium fimbriatum Wall., nom. nud. (Syntype K!), non Willd. 1810.
Rhizomes short, stout, erect, densely clothed with dull brown or red-brown, linear-lanceolate scales at apex. Fronds caespitose; fertile fronds (25--)60--80(--130) cm; stipes (10--)20--25(--40) cm long, (1--)2--3.5(--8) mm thick, bases densely clothed with the same scales as rhizomes, often pale red-purplish; laminae oblong or oblong-lanceolate, (15--)40--50(--80) cm long, (4--)15--30(--40) cm broad, long-acuminate at apex, bases roundish, pinnate, pinnae pinnatisect to 3-pinnate, ultimate pinnules pinnatilobate; pinnae 10--12(--20) pairs, alternate, upswept or slightly curved, petioled (5--8--10 mm), the lower 1--2 pairs a little shortened or almost not shortened, middle pinnae lanceolate, 15--30 cm long, 2--3(--10) cm broad, acuminate, caudate at apex, asymmetrical at base, the acroscopic side auricled and parallel to rach