Plants terrestrial, medium-sized. Rhizome erect or obliquely ascending, short, stout, rarely widely creeping, woody, dictyostelic, apex densely scaly; scales ferruginous, castaneous, brown, or black, glossy, ovate, broadly lanceolate, ovate-lanceolate, or lanceolate, entire or sparsely dentate or fimbriate, thick, consisting of slender, opaque cells with thick flexuose cell walls. Fronds caespitose or approximate, rarely remote, sometimes spirally arranged, segments catadromously or rarely anadromously arranged; stipe not articulate, with several separate vascular bundles, scaly, scales similar to those on rhizome, sometimes without scales on upper part of stipe; lamina broadly lanceolate, oblong, ovate, deltoid-ovate, or pentagonal, variously dissected, apex gradually reduced, rarely imparipinnate, once pinnate to quadripinnate or quinquepinnatifid; pattern of pinnation catadromous above base, commonly ± scaly, rarely glabrous; scales linear to lanceolate, bullate or flat, base cordate or truncate, apex subulate, entire or fimbriate on edges; rachis with longitudinal groove adaxially; ultimate segment base rounded and equilateral or rarely cuneate and inequilateral, margin usually serrate, rarely spinulose. Lamina papery or subleathery, rarely herbaceous, with or without hairs or setae adaxially, scaly, glabrous or with glands abaxially; costae and costules canaliculate adaxially and rising from rachis at ± acute angle; veins always free, pinnate or simple and undivided in once pinnate or bipinnatifid fronds, 2- or 3-forked in decompound fronds, all terminating in a prominent spindle-shaped hydathode short of frond margin. Sori orbicular, dorsal or rarely terminal on veins or veinlets, indusiate, rarely exindusiate; apophyses prominent or not; indusia persistent, superior or inferior, sessile or with a long stalk, orbicular-reniform to reniform or orbicular, rarely horseshoe-shaped, globose or subglobose, generally entire, smooth, rarely glandular or very rarely with erose margin, brownish, somewhat thick, sometimes thinly leathery, attached by a deep sinus on veins or veinlets. Spores monolete, surface verrucose or with broadly winged wall. n = 41.

The generic delimitation of Dryopteris adopted here follows the most recent molecular analysis, which discovered that Acrophorus, Acrorumohra, Diacalpe, Dryopsis, Nothoperanema, and Peranema were embedded within a paraphyletic Dryopteris and thus should all be merged into Dryopteris (Li Bing Zhang et al., BMC Evol. Biol. 12: 180. 2012). Most species of these genera share a short rhizome and catadromous arrangement of frond segments, unlike the sister genus of Dryopteris s.l., Arachniodes.

Excluded taxa

The following names are not treated due to lack of information: Dryopteris adaucta Rosenstock (Hedwigia 56: 341. 1915, type from Taiwan); D. camusiae Fraser-Jenkins (New Sp. Syndr. Indian Pteridol. 131. 1997, type from Yunnan); D. chimingiana Ching ex K. H. Shing & J. F. Cheng (Jiangxi Sci. 8(3): 47. 1990, type from Jiangxi); D. glabrior Ching & Z. Y. Liu (Bull. Bot. Res., Harbin 4(4): 11. 1984, type from Chongqing (Nanchuan), not Copeland (1910)); D. hezhangensis P. S. Wang (Pterid. Fl. Guizhou 315. 2001, type from Guizhou); D. jinfoshanensis Ching & Z. Y. Liu (Bull. Bot. Res. (Harbin) 6(1): 179. 1986; D. daozhenensis P. S. Wang & X. Y. Wang; D. nanchuanensis Ching & Z. Y. Liu (Jul 1984), not Ching & Z. Y. Liu (1983), nor Ching & Z. Y. Liu (Oct 1984); type from Chongqing (Nanchuan)); D. liyangensis Ching & Y. Z. Lan (Fl. Jiangsu. 1: 466. 1977, type from Jiangsu); D. lungnanensis Ching ex K. H. Shing & J. F. Cheng (Jiangxi Sci. 8(3): 48. 1990, type from Zhejiang); D. megacarpa Ching & Z. Y. Liu (Bull. Bot. Res., Harbin 4(4): 3. 1984, type from Chongqing (Nanchuan)); D. mollis (Swartz) Hieronymus var. subglabra Hosokawa (Trans. Nat. Hist. Soc. Taiwan 26: 78. 1936, type from Taiwan); D. parachinensis Ching & F. Z. Li (Bull. Bot. Res., Harbin 5(1): 157. 1985, type from Shandong); D. parachrysocoma Ching & Z. R. Wang (Acta Phytotax. Sin. 23(5): 344. 1985 ["para-chrysocoma"], type from Yunnan); D. peregrina C. Christensen (Index Filic. 284. 1905, based on Nephrodium regulare Baker, J. Bot. 13: 200-201. 1875, not Desvaux (1827), type

Shearer s.n. "China"); D. polylepis (Franchet & Savatier) C. Christensen var. pallida Ching (Bull. Fan Mem. Inst. Biol., Bot. 8: 422. 1938 ["pallidis"], type from Hubei); D. pseudocuspidata Christ (Bot. Gaz. 51: 357. 1911, type from Sichuan); D. shandongensis J. X. Li & F. Li (Acta Phytotax. Sin. 26: 406. 1988, type from Shandong); D. shanmenensis Ching & P. S. Chiu (Bot. Res. Acad. Sin. 2: 2. 1987, type from Guangxi); D. splendens (Hooker) Kuntze var. formosana Rosenstock (Hedwigia 56(5): 343. 1915, type from Taiwan); D. wenchuanensis H. S. Kung (Acta Bot. Yunnan. 4(4): 342. 1982, type from Sichuan); D. wuliangshanicola W. M. Chu ex S. G. Lu (Guihaia 11: 221. 1991, type from Yunnan); D. xiangxinensis S. F. Wu & C. M. Zhang (Keys Vasc. Pl. Wuling Mount. 570. 1995, type from Hunan); D. yenpingensis C. Christensen & Ching (Bull. Fan Mem. Inst. Biol., Bot. 8: 450. 1938, type from Fujian); D. zhenfengensis P. S. Wang & X. Y. Wang (Pterid. Fl. Guizhou, 337. 2001, type from Guizhou).

In addition, Dryopteris nigropaleacea (Fraser-Jenkins) Fraser-Jenkins (Bol. Soc. Brot., sér. 2, 55: 238. 1982; D. pallida Fomin subsp. nigropaleacea Fraser-Jenkins, Candollea 32(2): 316. 1977) and D. barbellata Fomin (Fl. Sibir. Orient. Extremi 5: 59. 1930), synonymized under D. sichotensis Komarov by Fraser-Jenkins (Bull. Brit. Mus. (Nat. Hist.), Bot. 14: 208 1986), have been recorded from S China and N China, respectively. These two species are not treated here due to insufficient material examined.

About 400 species: widely distributed in both hemispheres, mainly in Asia, especially from the Himalaya to China, Japan, and Korea; 167 species (60 endemic) in four subgenera in China.

(Authors: Wu Sugong (武素功), Xiang Jianying (向建英), Lu Shugang (陆树刚), Wang Faguo (王发国), Xing Fuwu (邢福武), Dong Shiyong (董仕勇), He Hai (何海), Zhang Libing (张丽兵); David S. Barrington, Maarten J. M. Christenhusz)