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.
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(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 spe