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FOC | Family List | FOC Vol. 2-3 | Dryopteridaceae

4. Dryopteris Adanson, Fam. Pl. 2: 20, 551. 1763.

鳞毛蕨属 lin mao jue shu

Authors: Sugong Wu, Xiang Jianying, Shugang Lu, Wang Faguo, Prof. Fuwu Xing, Shiyong Dong, He Hai, Li-Bing Zhang, David S. Barrington & Maarten J. M. Christenhusz

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.

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.

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.


1 (1) Lamina once imparipinnate, apical pinna similar to lower ones (1. D. subg. Pycnopteris)   (1)
+ Lamina once pinnate to quadripinnate or quinquepinnatifid, pinnae near apex gradually reduced to a pinnatifid apex   (2)
       
2 (1) Fronds with non-glandular hairs; indusia orbicular or reniform, inferior or superior (4. D. subg. Nothoperanema)   (3)
+ Fronds without hairs or with glandular or non-glandular hairs; indusia reniform, superior   (6)
       
3 (2) Indusia superior (above sori), flat, and reniform.   19 D. sect. Nothoperanema
+ Indusia inferior (beneath sori), globose or semiglobose   (4)
       
4 (3) Pinnae and pinnules of every order with or without a large cordate or ovate-lanceolate and often persistent scale at base; indusia membranous, semiglobose, indehiscent; lower portion of sporangiate stalk with a few multicellular clavate septate paraphyses; scales at base of stipe with multicellular clavate appendages on margin.   20 D. sect. Acrophorus
+ Pinnae and pinnules of every order without a large cordate scale at base; indusia leathery, globose, often splitting into 2 or 3 valves upon maturity; lower portion of sporangiate stalk with single-celled clavate septate paraphyses or without any paraphyses; scales at base of stipe with single-celled clavate appendages on margin or without any appendages   (5)
       
5 (4) Sori sessile; lower portion of sporangiate stalk with single-celled clavate septate paraphyses; scales at base of stipe with entire margins.   21 D. sect. Diacalpe
+ Sori stalked; lower portion of sporangiate stalk without any paraphyses; scales at base of stipe with single-celled clavate appendages on margin.   22 D. sect. Peranema
       
6 (2) Scales bullate (broad-based scales with ciliate apices) on stipe, rachis, and especially on costa and costule; if scales flat then grooves of pinna rachis and pinnule rachis closed near their bases and fronds with multicellular non-glandular hairs (D. sect. Dryopsis), or frond segments anadromously arranged (D. sect. Acrorumohra), or stipe and rachis scales stiff and narrowly lanceolate and filiform (D. liboensis) (3. D. subg. Erythrovariae)   (7)
+ Scales flat, not bullate on stipe and rachis (2. D. subg. Dryopteris)   (11)
       
7 (6) Frond segments anadromously arranged.   17 D. sect. Acrorumohra
+ Frond segments catadromously arranged   (8)
       
8 (7) Grooves of pinna rachis and pinnule rachis closed near their bases and fronds with multicellular non-glandular hairs.   18 D. sect. Dryopsis
+ Grooves of pinna rachis and pinnule rachis connected near their bases and fronds without hairs   (9)
       
9 (8) Lamina pentagonal-ovate, usually tripinnate; lowest basiscopic pinnule on lowest pinna markedly longer than those above; pinnule caudate and acutely pointed.   16 D. sect. Variae
+ Lamina lanceolate or ovate-lanceolate, usually pinnate to bipinnate; lowest basiscopic pinnule on lowest pinna not markedly longer than those above; pinnule not caudate, rounded   (10)
       
10 (9) Upper stipe with many small scales; scales on base of stipe lanceolate, brown or light brown.   14 D. sect. Erythrovariae
+ Upper stipe glabrous; scales on base of stipe linear-lanceolate, black or very dark brown.   15 D. sect. Indusiatae
       
11 (6) Segments asymmetrical at base   (12)
+ Segments symmetrical at base   (15)
       
12 (11) Lamina broadly lanceolate, basal pinnae symmetrical   (13)
+ Lamina triangular, basal pinnae asymmetrical, basiscopic basal pinnule proliferated   (14)
       
13 (12) Lamina tripinnatifid, with acute tooth on apex of segment.   10 D. sect. Remotae
+ Lamina bi- to quadripinnatifid, without acute tooth on apex of segment.   11 D. sect. Splendentes
       
14 (12) Lamina tri- to quadripinnate, stipe base scales filiform, golden.   12 D. sect. Purpurascentes
+ Lamina bi- to tripinnate, stipe base scales lanceolate or ovate-lanceolate, brown.   13 D. sect. Nephrocystis
       
15 (11) Lamina once pinnate, pinnae entire to deeply pinnatifid   (16)
+ Lamina twice pinnate or more dissected   (19)
       
16 (15) Pinnae entire to lobed, veins simple, basal veins of a segment (or at least acroscopic one) terminating ?halfway to segment margin.   1 D. sect. Hirtipedes
+ Pinnae pinnatifid or becoming nearly twice pinnate (at least several basal pairs), veins forked, rarely simple, terminating at segment margin   (17)
       
17 (16) Indusia membranous or strombuliform, ?thick, surrounding sporangia even when ripe.   2 D. sect. Pandae
+ Indusia not surrounding sporangia when ripe, sometimes deciduous   (18)
       
18 (17) Scales filiform or linear on rachis, costa, and abaxial surface of lamina.   3 D. sect. Fibrillosae
+ Scales lanceolate on rachis, costa, and abaxial surface of lamina.   4 D. sect. Dryopteris
       
19 (15) Lamina bipinnatifid, pinnules serrate   (20)
+ Lamina tripinnatifid or mostly tripinnatisect to quadripinnatifid   (21)
       
20 (19) Margin of pinnule or segment with long, acute teeth; indusia margin lacerate or erose.   5 D. sect. Caespitosae
+ Margin of pinnule or segment without long, acute teeth; indusia entire, cartilaginous.   6 D. sect. Pallidae
       
21 (19) Lamina oblong-lanceolate or broadly triangular, pinnae usually symmetrical.   7 D. sect. Marginatae
+ Lamina pentagonal, pinnae asymmetrical, basiscopic pinnule of pinnae longer   (22)
       
22 (21) Pinnae shortly stalked, ultimate segments with long, acute teeth.   8 D. sect. Lophodium
+ Pinnae long stalked, ultimate segments without long, acute teeth.   9 D. sect. Aemulae

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