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

4. Athyrium Roth, Tent. Fl. Germ. 3: 31, 58. 1799.

蹄盖蕨属 ti gai jue shu

Authors: Zhongren Wang & Masahiro Kato

Pseudocystopteris Ching.

Plants terrestrial, medium-sized. Rhizomes short, mostly erect, occasionally creeping or ascending, rarely long creeping; fronds caespitose, rarely approximate or distant. Stipe grooved abaxially, terete adaxially, often shortly glandular hairy in groove, base often swollen, with uniseriate tuberculate pneumathodes on lateral sides, narrowed toward very base, sometimes without pneumathodes, not narrowed to base, with vascular bundles 2, upward connected abaxially into U-shaped single bundle in section, scaly at base; scales red-brown, brown, or blackish brown, ovate-lanceolate, linear-subulate, or linear-lanceolate, entire, membranous, basifixed; lamina ovate, oblong, or broadly lanceolate, 1-3-pinnate; lamina segments inequilateral at base; groove of rachis and costae continuous to grooves of costae and costules, respectively; spines at base of costae, costules, and midribs on adaxial side present or absent, spines continuous from ridges on both sides of groove. Veins free, forked or pinnate, extending to tooth tips at lamina margin. Lamina usually herbaceous when dried, in a few cases papery or leathery, usually glabrate, rarely scaly or hairy, or shortly glandular hairy adaxially. Sori orbicular, orbicular-reniform, horseshoe-shaped, J-shaped, oblong, or shortly linear, abaxial, lateral to or crossing over veins; indusia brown, orbicular-reniform, horseshoe-shaped, J-shaped, lunate, oblong, or shortly linear, membranous, erose or ciliate, 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 or without folds. x = 40.

About 220 species: distributed mainly in the temperate zone and subtropical mountain forests; 123 species (probably more plus a number of hybrids) (69 endemic) in China.

The distribution center is located in the mountainous areas of SE China.

Athyrium is one of the largest genera of the pteridophytes and one of the taxonomically most difficult genera. Molecular data shows Athyrium as defined here is not monophyletic; Athyrium niponicum is transferred to Anisocampium, while Pseudocystopteris is nested within Athyrium. Hybridization makes the taxonomy of this genus more difficult. More than 60 hybrids have been known in Japan, and many additional hybrids are possible in China, requiring further analysis.

The following species, distributed in Taiwan, could not be treated here because no material was seen by the present authors: Athyrium puncticaule (Blume) T. Moore (Index Fil. 186. 1860; Aspidium puncticaule Blume, Enum. Pl. Javae 2: 159. 1828), Athyrium leiopodum (Hayata) Tagawa (Acta Phytotax. Geobot. 2(3): 195. 1933; Diplazium leiopodum Hayata, Icon. Pl. Formosan. 4: 217. 1914), and A. tenuicaule (Hayata) Tagawa (Acta. Phytotax. Geobot. 2(3): 195. 1933; Diplazium tenuicaule Hayata, Icon. Pl. Formosan. 4: 220. 1914), treated by Knapp as a synonym of A. leiopodum (Ferns Fern Allies Taiwan, 980. 2011). For more information, see Liu and Fraser-Jenkins (Taiwania 51: 293-297. 2006), Liu et al. (Fern Fl. Taiwan Athyrium, 26, 42. 2009), and Knapp (loc. cit.: 129-130, 439).

In addition, eleven uncertain taxa, not included in the following key, are listed at the end of the account.

Uncertain taxa

    Athyrium dulongicola W. M. Chu (Acta Bot. Yunnan., Suppl. 5: 41. 1992 [“dulongicolum”]), described from Yunnan. Athyrium excelsium Ching (Acta Bot. Boreal.-Occid. Sin. 6(1): 20. 1986), described from Yunnan. Athyrium fargesii Christ (Bull. Soc. Bot. France 52(Mém. 1): 49. 1905), described from Yunnan. Athyrium filix-femina (Linnaeus) Roth subvar. brevidens Christ (Bull. Soc. Bot. France 52(Mém. 1): 46. 1905), described from Yunnan. Athyrium filix-femina (Linnaeus) Roth var. duclouxii Christ (Bull. Soc. Bot. France 52(Mém. 1): 46. 1905), described from Yunnan. Athyrium filix-femina (Linnaeus) Roth var. paleosum Christ (Notul. Syst. (Paris) 1: 48. 1909), described from Yunnan. Athyrium ×heterosporum Y. T. Hsieh & Z. R. Wang (Acta Phytotax. Sin. 27: 156. 1989), described from Yunnan and treated in FRPS (3(2): 266. 1999). Athyrium nemotum Ching (Acta Bot. Boreal.-Occid. Sin. 6(3):149-150. 1986), described from Guangxi. Athyrium nigripes (Blume) T. Moore var. elongatum Christ (Bull. Herb. Boissier 7(1): 13. 1899), described from Yunnan. Athyrium rotundifolium Ching (Fl. Fujian. 1: 597. Apr 1982), described from Fujian. Athyrium silvestrii Christ (Nuovo Giorn. Bot. Ital. n.s., 17. 226. 1910), described from Hubei.


1 Sori small, orbicular-reniform   (2)
+ Sori prominent, elongate, horseshoe-shaped, or J-shaped   (8)
       
2 (1) Plants medium-sized to large; fronds usually more than 60-70 cm, up to 2-3 m, all pinnae stalked.   117 A. atkinsonii
+ Plants small to medium-sized; fronds usually less than 60-70 cm, second basal pair of pinnae and all upper pinnae subsessile or very shortly stalked   (3)
       
3 (2) Lamina triangular; basal pinnae largest, narrowly elliptic to elliptic-oblanceolate with lower pinnules gradually shorter to base, prominently narrowed at base, pinnules of upper pinnae often catadromous; mostly in mountains below 3000 m   (4)
+ Lamina ovate to oblanceolate; basal pinnae ?shorter than upper, ovate to oblong-lanceolate, rarely subauriculate, not gradually shortened toward base, at most 1 or 2 basal pairs of pinnules slightly shortened, pinnules or segments of all pinnae anadromous or subopposite   (5)
       
4 (3) Rachis and costae glabrate abaxially, pinnules ?rounded at base, distant from costae, teeth at segment margin long, pointed.   118 A. spinulosum
+ Rachis and costae ?glandular hairy abaxially, pinnules subtruncate at base, often close to costae, teeth short and narrow.   119 A. subtriangulare
       
5 (3) Lamina ovate to ovate-oblong, usually 2- or 3-pinnate; pinnules inequilateral at base, shortly stalked; rachis and costae without long multicellular hairs   (6)
+ Lamina oblanceolate or ovate-oblong, 2-pinnatifid to 2-pinnate; segments adnate to costae by narrow wing, if bipinnate then pinnules subequilateral at base, sessile; rachis, costae, and veins with sparse long multicellular hairs   (7)
       
6 (5) Fronds less than 35 cm; ultimate segments approximate, teeth at segment margin short.   120 A. schizochlamys
+ Fronds up to 55 cm; ultimate segments separate, teeth at segment margin long, thin.   121 A. atuntzeense
       
7 (5) Lamina oblanceolate, many pairs of lower pinnae gradually shorter than upper pinnae.   122 A. davidii
+ Lamina ovate-oblong, basal 2 pairs of pinnae slightly shortened.   123 A. repens
       
8 (1) Sori orbicular or elliptic, exindusiate or indusia caducous; perispore with folds   (9)
+ Sori elliptic, shortly linear, J-shaped, horseshoe-shaped, or reniform; indusia persistent, rarely incomplete but visible, at least when young; perispore with or without folds   (11)
       
9 (8) Stipe and rachis dark brown, throughout with dense castaneous-black or dark brown, ovate-lanceolate scales and fibrous scales; basal pinnae tapering to base; pinna apex obtuse; stipe much shorter than lamina.   1 A. wallichianum
+ Stipe and rachis stramineous or pale purplish red, scales dense, brown at base; basal pairs of pinnae slightly abbreviated; stipe slightly shorter than lamina; pinna apex acuminate   (10)
       
10 (9) Lamina 1-pinnate, pinnae pinnatilobate to pinnatipartite, occasionally 2-pinnate; pinnae sessile or subsessile (stalk less than 3 mm); teeth at lobe margin short and pointing outward.   2 A. dissitifolium
+ Lamina 2-pinnate; pinnae prominently stalked (stalk up to 10 mm or more); teeth at lobe margin long and ascending.   3 A. exindusiatum
       
11 (8) Rhizomes long or shortly creeping with fronds distant or approximate, or erect   (12)
+ Rhizomes erect or ascending; fronds caespitose   (23)
       
12 (11) Rhizomes long creeping; costae and costules with prominent spines at base on adaxial side   (13)
+ Rhizomes creeping or erect; costae and costules without spines at base adaxially; perispore with folds   (14)
       
13 (12) Plants medium-sized, more than 20 cm tall; lamina 3-pinnate or more divided; sori not close to costae or costules, indusia not overlaying midribs and veins; perispore with folds.   4 A. pectinatum
+ Plants small, less than 20 cm tall; lamina 1-pinnate, pinnae pinnatipartite; sori close to costae or costules, indusia often overlaying costae or costules; perispore without folds.   99 A. chingianum
       
14 (12) Rhizomes long creeping; fronds distant   (15)
+ Rhizomes shortly creeping (occasionally long creeping); fronds approximate, or rhizome erect   (17)
       
15 (14) Lamina subdimorphic, stipe of sterile frond shorter than lamina, but stipe of fertile frond much longer than lamina; lamina deltoid-ovate; pinnae 3-5 pairs; basal pinnae not abbreviated.   5 A. ruilicola
+ Lamina monomorphic, stipe shorter than lamina or equal; lamina ovate-oblong; pinnae 15-20 pairs; basal pair of pinnae abbreviated   (16)
       
16 (15) Lamina 2-pinnate, adaxial surface glabrate; teeth not long apiculate.   6 A. biserrulatum
+ Lamina 3-pinnate, adaxial surface pubescent; teeth long apiculate.   7 A. wumonshanicum
       
17 (14) Lamina ovate to narrowly ovate, apex cuspidate; pinnae stalked; stipe only slightly shorter than lamina; scales at stipe base pale brown; sori shortly linear, oblong, or narrowly J-shaped   (18)
+ Lamina lanceolate, oblong-lanceolate, or oblanceolate, apex acuminate; pinnae sessile or occasionally shortly stalked (stalk up to 5 mm); stipe much shorter than lamina; scales at stipe base red-brown or castaneous; sori suborbicular, elliptic, reniform, or shortly J-shaped   (19)
       
18 (18) Pinnae usually 1-pinnate or more dissected; pinnules ovate, oblong-lanceolate, or falcate-lanceolate, shallowly lobed to pinnatipartite, basal pinnules usually shortly stalked.   A. niponicum (see Anisocampium niponicum, p. 448)
+ Pinnae usually pinnatipartite to pinnatisect; lobes falcate-lanceolate, at most shallowly lobed, decurrent to broad wings of costa on basiscopic side.   8 A. brevisorum
       
19 (17) Indusia pale brown, entire; basal pinnae not shortened or nearly so; rhizome erect   (20)
+ Indusia pallid, margin ciliate; many basal pinnae gradually shortened to base or only 1 or 2 pairs of basal pinnae prominently shortened; rhizome shortly creeping   (21)
       
20 (19) Scales at stipe base yellow-brown; lamina pinnate with pinnae pinnatipartite or 2-pinnate with pinnules pinnatilobate; pinnae narrowly lanceolate.   9 A. yokoscense
+ Scales at stipe base red-brown, central part black-brown; lamina pinnate, pinnae pinnatilobate to pinnatipartite, broadly ovate.   10 A. fauriei
       
21 (19) Teeth at lamina margin obtuse.   11 A. fallaciosum
+ Teeth at lamina margin sharp and long   (22)
       
22 (21) Rachis and costae glabrate abaxially.   12 A. rupicola
+ Rachis and costae pubescent abaxially.   13 A. glandulosum
       
23 (11) Costae (or costules and midribs) without spines at base on adaxial side, at most larger plants rarely with extremely short processes in distal part of costa; perispore with or without folds   (24)
+ Costae (or also costules and midribs) with long or short spines at base on adaxial side; perispore without folds   (48)
       
24 (23) Pinnae prominently broadened at base, inequilateral, auriculate at acroscopic base, usually cuneate basiscopically; in pinnate or more divided lamina, all lobes or pinnules anadromous; costae without spines on adaxial side; perispore with folds (rarely without folds)   (25)
+ Pinnae narrowed at base, not or slightly broadened, equilateral or subequilateral; at least pinnules or lobes of upper pinnae catadromous or opposite; costae of larger plants rarely with very short processes adaxially; perispore without folds   (33)
       
25 (24) Fertile fronds less than 10 cm; perispore without folds.   14 A. minimum
+ Fertile fronds more than 10 cm; perispore with folds   (26)
       
26 (25) Lamina pinnate, pinnae repand or shallowly deltoid-lobed, occasionally acroscopic basal lobes of basal pinnae pinnatisect; rachis glandular pubescent   (27)
+ Lamina pinnate to 2-pinnate or more divided, if pinnate then lobes of pinnae not deltoid; rachis without glandular hairs (occasionally with pale brown, short unicellular hairs)   (28)
       
27 (26) Pinna obtuse at apex, basal acroscopic auricles entire or slightly repand to dentate, rachis with sparse black or brown glandular hairs.   15 A. nakanoi
+ Pinna acute at apex, basal acroscopic auricles denticulate at apex, rachis with sparse brown glandular hairs.   16 A. zhenfengense
       
28 (26) Lamina ovate to ovate-triangular, basal pinnae largest, narrowly ovate, basal 1 or 2 pairs of pinnules of basal pinnae slightly shortened; ultimate pinnules or lobes obtuse.   17 A. bucahwangense
+ Lamina narrowly lanceolate to ovate-lanceolate, basal pinnae slightly shortened, deltoid-ovate to deltoid-lanceolate, basal pair of pinnules of basal pinnae largest; ultimate pinnules or lobes acute or rounded   (29)
       
29 (28) Rachis without glandular hairs on adaxial side.   18 A. erythropodum
+ Rachis glandular pubescent adaxially   (30)
       
30 (29) Lamina thickly papery or thinly leathery, dark green, surface shiny; indusia small, partly fugacious when mature.   19 A. drepanopterum
+ Lamina papery or herbaceous, green, surface not shiny; indusia prominent, persistent   (31)
       
31 (30) Lamina 2- or 3-pinnate; stipe pale purplish red, scales at base reddish brown; pinnae and ultimate pinnules acute at apex; sori less than 1 mm in diam..   20 A. fimbriatum
+ Lamina 1- or 2-pinnate; stipe stramineous, scales at base brown or dark brown; pinnae and ultimate lobes rounded at apex; sori 1-2 mm in diam   (32)
       
32 (31) Lamina usually lanceolate, pinnate, at most 2-pinnate at base, but free pinnules not more than 5 pairs.   21 A. anisopterum
+ Lamina usually ovate-lanceolate, 2-pinnate, free pinnules of middle pinnae more than 10 pairs.   22 A. foliolosum
       
33 (24) Lamina densely pubescent on adaxial surface   (34)
+ Lamina glabrate   (35)
       
34 (33) Rachis and costae pubescent on abaxial side; pinna lobes obtuse at apex.   23 A. sericellum
+ Rachis and costae glabrate on abaxial side; pinna lobes rounded at apex.   24 A. suprapubescens
       
35 (33) Basal pinnae prominently constricted at base; sori mostly shortly linear; rachis and costae glabrate or with short straight glandular hairs abaxially   (36)
+ Basal pinnae slightly broadened or slightly constricted at base; sori mostly reniform, horseshoe-shaped, J-shaped, or oblong; rachis and costae with sparse squarrose-glandular hairs abaxially   (38)
       
36 (35) Lamina deltoid-ovate, apex cuspidate; basal several pairs of pinnae subopposite, basiscopic lobes of basal pinnae longer than acroscopic ones.   25 A. hainanense
+ Lamina lanceolate or ovate-oblong, apex acuminate; pinnae alternate, basiscopic lobes of basal pinnae nearly as long as acroscopic ones   (37)
       
37 (36) Indusia poorly developed, visible only when young; teeth at lobe margin cuspidate.   26 A. pachyphyllum
+ Indusia well developed, persistent; teeth at lobe margin not cuspidate.   27 A. wangii
       
38 (35) Lamina broadly ovate or ovate, basal 1 or 2 pairs of pinnae slightly shortened or not shortened   (39)
+ Lamina narrowly ovate, elliptic-lanceolate, or oblanceolate, more than 2 or 3 pairs of basal pinnae gradually shortened   (41)
       
39 (38) Scales at stipe base black, shiny.   28 A. melanolepis
+ Scales at stipe base brown or dark brown, not shiny   (40)
       
40 (39) Plants up to 120 cm; lamina herbaceous, stipe and rachis stramineous or pale purplish red, not shiny.   29 A. brevifrons
+ Plants to 65 cm; lamina thinly herbaceous, stipe and rachis pale reddish, shiny.   30 A. rubripes
       
41 (38) Lamina narrowly ovate, basal 2 or 3 pairs of pinnae shortened, but basal pinnae not shortened as auricles.   31 A. sinense
+ Lamina elliptic-lanceolate or oblanceolate, more than 4 pairs of basal pinnae gradually shortened, basal pinnae often shortened as auricles   (42)
       
42 (41) Scales at stipe base lanceolate or linear-lanceolate, less than 1.5 mm wide; lamina pinnate, pinnae pinnatipartite, lobes serrate, lateral veins usually simple.   32 A. attenuatum
+ Scales at stipe base ovate or deltoid-ovate, more than 3-5 mm wide; lamina usually 2-pinnate, pinnules shallowly lobed or biserrate, lateral veins usually 2-4-furcate   (43)
       
43 (42) Lamina black when dried; scales at stipe base castaneous at middle, brown in marginal part; pinnule rounded-truncate at apex.   33 A. tarulakaense
+ Lamina not black when dried; scales at stipe base brown or dark brown; pinnule obtuse at apex   (44)
       
44 (43) Pinnae deltoid-ovate or deltoid-lanceolate, less than 3-4 cm, approximate.   34 A. contingens
+ Pinnae linear-lanceolate, more than 4 cm, spaced apart from each other   (45)
       
45 (44) Pinnae shortly stalked, slightly constricted toward base, pinnules up to 2.5-3 cm.   35 A. nudifrons
+ Pinnae subsessile, not constricted toward base, pinnules usually less than 2.5-3 cm   (46)
       
46 (45) Middle pinnae ca. 1 cm wide, separated from each other by space more than width of pinna.   36 A. nephrodioides
+ Middle pinnae more than 1 cm wide, separated from each other by space much less than width of pinna   (47)
       
47 (46) Pinnae 3-6 ?less than 1.8 cm, prominently ascending.   37 A. adscendens
+ Pinnae up to 16-18 ?up to 3 cm, spreading or slightly ascending.   38 A. dentigerum
       
48 (23) Lamina pinnate, oblanceolate, pinnae pinnatilobate to pinnatipartite, many pairs of basal pinnae gradually shortened, basal pinnae often auricles.   39 A. mehrae
+ Lamina 1-3-pinnate, not oblanceolate, basal pinnae not shortened, or at most 1 to several pairs slightly shortened, basal pinnae never auricles   (49)
       
49 (48) Lamina narrowly lanceolate, pinnae more than 20-25 pairs   (50)
+ Lamina broader, not narrowly lanceolate, pinnae less than 20-25 pairs   (51)
       
50 (49) Lamina usually less than 35 cm, 1- or 2-pinnate, lobe margin dentate, costae with short spines adaxially.   40 A. multipinnum
+ Lamina up to 35 cm or more, 2-pinnate, lobe margin long serrulate, costae with long spines adaxially.   41 A. elongatum
       
51 (49) Indusia various, J-shaped, horseshoe-shaped, reniform, elliptic, or shortly linear; scales at stipe base often yellow-brown, brown, or dark brown   (52)
+ Indusia usually shortly linear or oblong; scales at stipe base often black or dark brown   (83)
       
52 (51) Pinnules or lobes of upper pinnae anadromous, occasionally catadromous or subopposite; rachis and costae stramineous, rarely pale purplish red, glabrate or sparsely pubescent on abaxial side; margin of narrow wings along costae or sinus between lobes glabrate   (53)
+ Pinnules or lobes of upper pinnae catadromous or subopposite; rachis and costae usually pale purplish red, occasionally stramineous, pubescent on abaxial side; margin of narrow wings of costa or sinus between lobes sparsely pubescent or glabrate   (61)
       
53 (52) Pinnules or lobes of upper pinnae opposite or subopposite   (54)
+ Pinnules or lobes of upper pinnae anadromous, alternate, or only basal pair subopposite   (55)
       
54 (53) Pinnules or lobes downward reflexed or subspreading.   42 A. oppositipennum
+ Pinnules or lobes spreading.   43 A. crassipes
       
55 (53) Pinnae (especially also lamina tip) or pinnules ascending or at most subspreading   (56)
+ Pinnae (especially also lamina tip) or pinnules downward reflexed   (58)
       
56 (55) Lower pinnae alternate, at most basal pinnae subopposite; rachis and costae pale purplish red, spines prominent on adaxial side, but costules without spines.   44 A. tripinnatum
+ Basal pinnae to several pairs of lower pinnae opposite or subopposite; rachis and costae stramineous, with very short spines adaxially, costules also with spines   (57)
       
57 (56) Basal pinnules subopposite, rachis and costae sparsely pubescent abaxially.   45 A. deltoidofrons
+ Basal pinnules anadromous, rachis and costae glabrate abaxially.   46 A. delicatulum
       
58 (55) Lamina pinnate, pinnae at most pinnatipartite, sessile, downward reflexed, upper pinnae spreading.   47 A. reflexipinnum
+ Lamina 1- or 2-pinnate, pinnae shortly stalked, usually spreading or ascending, distal pinnae downward reflexed; pinnules pinnatisect, reflexed downward or spreading   (59)
       
59 (58) Sori horseshoe-shaped; pinnules lanceolate, subsessile.   48 A. devolii
+ Sori oblong or J-shaped; pinnules ovate-deltoid or oblong, prominently stalked or adnate to narrow wing of costa   (60)
       
60 (59) Pinnules ovate-deltoid, acute at apex, prominently stalked (stalk ca. 1 mm).   49 A. baishanzuense
+ Pinnules oblong, rounded-obtuse at apex, adnate to narrow wing of costa.   50 A. maoshanense
       
61 (52) Indusia usually reniform, orbicular-reniform, or elliptic; lamina broadly ovate, apex acuminate, basal pinnae largest; scales yellow-brown.   51 A. omeiense
+ Indusia usually oblong, J-shaped, or horseshoe-shaped; lamina usually oblong-ovate, apex cuspidate or acuminate, basal pinnae subequal to second basal pinnae or slightly larger; scales brown or dark brown   (62)
       
62 (61) Pinnae prominently stalked, stalk usually more than 2 mm, alternate, basal pinnae subopposite; margin of narrow wings of costa or sinus between lobes glabrate or sparsely pubescent; indusia subentire or erose   (63)
+ Pinnae usually sessile, occasionally shortly stalked, stalk less than 2 mm, basal 3 or more pairs of pinnae often opposite or subopposite; margin of narrow wings of costa or sinus between lobes usually sparsely pubescent, occasionally glabrate; indusia usually ciliate, occasionally lacerate or erose   (69)
       
63 (62) Margin of narrow wings of costae sparsely pubescent, costae densely pubescent abaxially   (64)
+ Margin of narrow wings of costae glabrate, rachis and costae glabrate or sparsely pubescent   (65)
       
64 (63) Adaxial surface of lamina and lobe margin sparsely pubescent; pinnules narrowly deltoid to deltoid-oblong, acute at apex.   57 A. ludingense
+ Adaxial surface of lamina and lobe margin glabrate; pinnules oblong to deltoid-ovate, obtuse at apex.   58 A. hirtirachis
       
65 (63) Pinnules oblong, rounded-obtuse at apex.   52 A. intermixtum
+ Pinnules ovate to oblong-lanceolate, acute at apex   (66)
       
66 (65) Lamina apex usually acuminate; rachis and costae glabrate on abaxial side; pinnules of middle and basal pinnae pinnatipartite to pinnatisect   (67)
+ Lamina apex often cuspidate; rachis and costae sparsely pubescent abaxially; pinnules of middle and basal pinnae pinnatilobate to pinnatipartite   (68)
       
67 (66) Scales pale brown, linear-lanceolate; costae pale purplish red on abaxial side.   53 A. costulalisorum
+ Scales dark brown, ovate to ovate-lanceolate; costae stramineous.   54 A. dentilobum
       
68 (66) Costae pale purplish red; lamina thinly herbaceous, apex cuspidate prominently.   55 A. vidalii
+ Costae stramineous; lamina papery, apex cuspidate often not prominently.   56 A. mackinnonii
       
69 (62) Rachis and costae stramineous, with dense unicellular straight hairs on abaxial side, with very short spines on adaxial side; midribs without spines.   59 A. himalaicum
+ Rachis and costae usually pale purplish red, occasionally stramineous, sparsely pubescent or glabrate abaxially, with long or short spines adaxially; costules or midribs with or without spines   (70)
       
70 (69) Spines on adaxial surface of costae very short, but if long then costules or midribs without spines   (71)
+ Spines on adaxial surface of costae long, narrowly deltoid; if lamina 2-pinnate or more dissected then costules or midribs also spinose   (76)
       
71 (70) Plants less than 30 cm tall; pinnae less than 5 cm, pinnatipartite; rachis and costae pale purplish red or stramineous; margin of narrow wings of costae sparsely pubescent   (72)
+ Plants more than 30 cm tall; pinnae more than 7 cm, pinnate; rachis and costae stramineous; margin of narrow wings of costae sparsely pubescent or glabrate   (73)
       
72 (71) Basal pinnae prominently shortened; spines on adaxial surface of costae very short; rachis and costae stramineous; adaxial surface of lamina glabrate.   60 A. emeicola
+ Basal pinnae not shortened; spines on adaxial surface of costae sharp and long; rachis brown-stramineous, costae pale purplish red abaxially; adaxial surface of lamina with sparse pallid short hairs.   61 A. bomicola
       
73 (71) Spines on adaxial surface of costae sharp and long; margin of narrow wings of costae glabrate.   62 A. flabellulatum
+ Spines on adaxial surface of costae very short; margin of narrow wings of costae sparsely hairy   (74)
       
74 (73) Plants less than 50 cm tall; pinnules oblong, ca. 7 mm, apex rounded, margin crenate.   63 A. daxianglingense
+ Plants to 1 m tall; pinnules broadly lanceolate, 22-27 mm, apex obtuse, margin incised   (75)
       
75 (74) Middle pinnae more than 20 cm; pinnules more than 20 pairs, oblong-lanceolate or oblong-ovate.   64 A. obtusilimbum
+ Middle pinnae less than 12 cm; pinnules ca. 14 pairs, deltoid-ovate or broadly deltoid-lanceolate.   65 A. baoxingense
       
76 (70) Margin of narrow wings of costae and rachis glabrate   (77)
+ Margin of narrow wings of costae and rachis ?with unicellular short hairs   (78)
       
77 (76) Lamina thinly herbaceous, teeth at lobe margin long and ascending, sori 2-4 pairs per pinnule.   66 A. jinshajiangense
+ Lamina papery or firm-herbaceous, lobe margin dentate, sori 4-7 pairs per pinnule.   67 A. interjectum
       
78 (76) Adaxial surface of lamina with unicellular short hairs.   68 A. suprapuberulum
+ Adaxial surface of lamina glabrate   (79)
       
79 (78) Pinnae less than 1 cm wide, apex rounded-truncate; scales at stipe base black in center, sharply with narrow pale brown margin.   69 A. zayuense
+ Pinnae usually more than 2 cm wide, apex obtuse; scales at stipe base dark brown in center, gradually paler toward margin   (80)
       
80 (79) Basal pinnules often reflexed.   70 A. lineare
+ Pinnules spreading or ascending   (81)
       
81 (80) Lamina 2-pinnate with pinnules pinnatisect to 3-pinnate with secondary pinnules pinnatilobate; basal pinnae much larger than upper pinnae, oblong-ovate; costae and costules with short spines on adaxial side.   71 A. chungtienense
+ Lamina 1-pinnate with pinnae pinnatisect to 2-pinnate with pinnules pinnatilobate; basal pinnae equal to upper pinnae or slightly shorter, narrowly lanceolate or narrowly oblanceolate; only in large pinnae costules with spines adaxially   (82)
       
82 (81) Base of middle pinnae inequilateral; teeth at pinnule margin or lobe margin sharp and long.   72 A. austro-orientale
+ Base of middle pinnae subequilateral; teeth at pinnule margin or lobe margin short.   73 A. dubium
       
83 (51) Costae and costules with long spines adaxially, ultimate midribs often also with spines   (84)
+ Costae with subulate, short spines on adaxial side; costules or midribs without spines, occasionally with very short spines   (104)
       
84 (83) Lamina 3-pinnate or more divided; secondary pinnules prominently stalked; rachis and costae glabrate abaxially   (85)
+ Lamina 1- or 2-pinnate with pinnules at most pinnatifid; rachis and costae pubescent or glabrate abaxially   (87)
       
85 (84) Lamina deltoid or ovate-deltoid; ultimate lobes broad, suboblong, usually more than 2-3 mm wide.   96 A. adpressum
+ Lamina elliptic; ultimate lobes slender and long, linear or narrowly subulate, usually ca. 1 mm wide   (86)
       
86 (85) Ultimate lobes linear, ascending; sori costular, straight.   97 A. rhachidosorum
+ Ultimate lobes narrowly subulate, spreading; sori subcostular, often curved.   98 A. araiostegioides
       
87 (84) Sori oblong or shortly linear, straight, close to midribs, often partly overlaying them when mature; rachis and costae glabrate abaxially, at most costae slightly pubescent at base   (88)
+ Sori curved or straight, not close to midribs, not overlaying them when mature; rachis and costae pubescent abaxially, rarely glabrate   (94)
       
88 (87) Fronds less than 25 cm; lamina narrowly deltoid or deltoid-ovate, glabrous or glandular hairy on adaxial surface   (89)
+ Fronds more than 30 cm; lamina broadly ovate to linear-lanceolate; lamina adaxial surface glabrate   (91)
       
89 (88) Rhizome creeping; lamina glabrous adaxially.   99 A. chingianum
+ Rhizome erect; lamina glandular hairy adaxially   (90)
       
90 (89) Lamina deltoid, basal pinnae not shortened; pinna lobes subentire; sori close to costa.   100 A. supraspinescens
+ Lamina ovate-deltoid, basal pinnae slightly shortened; pinnules crenate or serrate-lobed; sori distant from costa.   101 A. medogense
       
91 (88) Lamina pinnate; pinnae deltoid-lanceolate, less than 3.5 cm, at most pinnatisect, pinna lobes approximate.   102 A. yui
+ Lamina 2-pinnate; pinnae lanceolate or linear-lanceolate, 5-20 cm, pinnules separate from each other   (92)
       
92 (91) Lamina ovate to oblong-lanceolate; pinnule lobes subentire or only slightly serrulate at margin.   103 A. fangii
+ Lamina broadly ovate to ovate; pinnule lobes with prominently long or short teeth at margin   (93)
       
93 (92) Pinnae 13-16 pairs; pinnules oblong; rachis and costae glabrate abaxially.   104 A. nyalamense
+ Pinnae 6 or 7 pairs; pinnules obovate or oblong; rachis and costae densely pubescent abaxially.   105 A. yuanyangense
       
94 (87) Lamina lanceolate to linear-lanceolate; pinnae usually more than 18 pairs; rachis usually with bulbils below apex, occasionally without bulbils   (95)
+ Lamina broadly ovate, ovate, or lanceolate; pinnae less than 15 pairs; rachis usually without bulbils below apex, occasionally bulbiferous   (97)
       
95 (94) Lamina narrowly lanceolate to linear-lanceolate; many basal pinnae shortened; pinnae 3-4 cm, subsessile; pinna lobes less than 10 pairs, minutely toothed at margin; bulbils usually large.   106 A. clarkei
+ Lamina lanceolate to narrowly lanceolate; only 1 or 2 pairs of basal pinnae slightly shortened; pinnae 6-11 cm, usually shortly stalked; pinna lobes more than 10 pairs, biserrate or lobed at margin; bulbils small   (96)
       
96 (95) Pinnules deltoid-broadly lanceolate; lobes minutely toothed at margin.   107 A. viviparum
+ Pinnules ovate-oblong; lobes prominently toothed, teeth ca. 1 mm.   108 A. strigillosum
       
97 (94) Plants less than 30 cm tall; lamina lanceolate, less than 6 cm wide; pinnae narrow and small, less than 1 cm wide; sori close to midribs.   109 A. nigripes
+ Plants usually more than 30 cm tall; lamina broadly ovate or ovate, more than 6 cm wide; pinnae more than 1 cm wide; sori medial or inframedial   (98)
       
98 (97) Pinnules pinnatilobate to pinnatifid, rarely pinnatipartite, apex obtuse, margin crenate; sori straight; lamina papery when dry   (99)
+ Pinnules pinnatifid to pinnatisect, apex acute or obtuse, margin with sharp teeth; sori curved and straight or straight; lamina herbaceous when dried   (100)
       
99 (98) Lamina oblong-ovate; basal several pairs of pinnae opposite or subopposite; stipe and rachis stramineous.   110 A. guangnanense
+ Lamina subdeltoid; basal pinnae subopposite, most other pinnae alternate; stipe and rachis usually pale purplish red.   111 A. silvicola
       
100 (98) Lamina ovate to narrowly ovate, occasionally lanceolate; pinnules or lobes separate from each other; pinnules oblong-ovate, inequilateral; sori straight, curved or variously shaped; rachis often bearing bulbils below apex   (101)
+ Lamina broadly ovate or ovate; pinnules or lobes approximate; pinnules oblong-ovate to narrowly deltoid-ovate, subequilateral; sori straight, rarely curved; rachis without bulbils below apex   (102)
       
101 (100) Pinnules of middle pinnae anadromous.   112 A. cryptogrammoides
+ Pinnules of middle pinnae subopposite.   113 A. iseanum
       
102 (100) Stipe and rachis pale purplish red, fragile; costae densely pubescent on abaxial side.   114 A. imbricatum
+ Stipe and rachis stramineous, tough; costae glabrate abaxially   (103)
       
103 (102) Stipe usually longer than lamina; lamina ovate, apex cuspidate; pinnae less than 10 pairs.   115 A. xichouense
+ Stipe usually shorter than lamina; lamina oblong, apex acuminate; pinnae ca. 13 pairs.   116 A. nanyueense
       
104 (83) Lamina broadly ovate or ovate, apex often cuspidate, occasionally narrowly ovate, apex shortly acuminate; pinnae usually prominently stalked (stalk usually more than 2-3 mm)   (105)
+ Lamina oblong-ovate or lanceolate, occasionally ovate, acuminate at apex, occasionally cuspidate; pinnae usually sessile, or shortly stalked (stalk less than 2 mm)   (112)
       
105 (104) Pinnules of basal 2 or 3 pairs of pinnae anadromous, pinnae prominently stalked (stalk 3-5 mm); rachis and costae glabrate on abaxial side.   74 A. clivicola
+ Pinnules of basal pinnae anadromous, pinnules of other pinnae subopposite or catadromous, pinnae shortly stalked; rachis and costae pubescent on abaxial side, rarely glabrate   (106)
       
106 (105) Pinnules shallowly lobed, lobes up to 1 mm, or bidentate.   75 A. infrapuberulum
+ Pinnules subentire, or serrate or lobed   (107)
       
107 (106) Pinnules lanceolate to narrowly lanceolate, 3 or more ?as long as wide; sori subparallel to midrib; indusia erose.   76 A. criticum
+ Pinnules deltoid, oblong-ovate, or ovate, occasionally narrowly ovate, less than 3 ?as long as wide; sori not parallel to midrib; indusia entire or subentire   (108)
       
108 (107) Lamina narrowly deltoid-ovate, narrowly ovate or ovate, pinnae 7 pairs or more except pinnae of cuspidate apex; pinnules narrow, usually less than 5 mm wide at middle, subentire; pinna rachis sparsely pubescent on abaxial side   (109)
+ Lamina deltoid or deltoid-ovate, pinnae ca. 5 pairs below cuspidate apex; pinnules usually more than 5 mm wide at middle, serrate; costae densely pubescent on abaxial side   (110)
       
109 (108) Lamina ovate-oblong, cuspidate at apex; pinnae up to 18 cm, pinnules more than 10 pairs.   77 A. brevistipes
+ Lamina narrowly deltoid-ovate, acuminate at apex; pinnae 5-11 cm, pinnules less than 10 pairs.   78 A. kenzo-satakei
       
110 (108) Base of basal pinnae broadened with basal pinnules largest; pinnules auriculate at base on both sides.   79 A. longius
+ Lower pinnules of basal pinnae shortened; pinnules auriculate only at acroscopic base   (111)
       
111 (110) Pinnules of middle pinnae ascending, oblong, serrulate at margin.   80 A. wardii
+ Pinnules of middle pinnae broadly deltoid-lanceolate, shallowly lobed to pinnatifid.   81 A. venulosum
       
112 (104) Pinnules or pinnule lobes rounded-obtuse or truncate at apex, margin rounded lobed or crenate; sori usually medial.   82 A. christensenii
+ Pinnules or pinnule lobes acute or obtuse at apex, lobed or toothed at margin, not crenate; sori usually close to costule   (113)
       
113 (112) Basal pinnules overlaying rachis   (114)
+ Basal pinnules not overlaying rachis   (116)
       
114 (113) Pinnae acuminate at apex; pinnules obtuse at apex, teeth at margin spreading.   83 A. delavayi
+ Pinnae caudate-acuminate at apex; pinnules acute or obtuse at apex, teeth at margin oblique   (115)
       
115 (114) Pinnae 14-16 cm, caudate or long acuminate at apex; pinnules shortly acuminate, long auriculate at acroscopic base, midrib without spines on adaxial side.   84 A. caudiforme
+ Pinnae 10-12 cm, shortly caudate at apex; pinnules acute, shortly auriculate at acroscopic base, midrib with spines on adaxial side.   85 A. subrigescens
       
116 (113) Scales at stipe base ovate-lanceolate or lanceolate; pinnules of upper pinnae anadromous or subopposite; sori ca. 1 mm, close to midrib, almost parallel to midrib.   86 A. decorum
+ Scales at stipe base linear to linear-lanceolate; pinnules of upper pinnae catadromous or subopposite; sori more than 1.5 mm, ascending to midrib   (117)
       
117 (116) Pinnae often shortly stalked; pinnules narrowly oblong, broadly cuneate at base; costae and distal part of rachis with very short spines adaxially.   87 A. uniforme
+ Pinnae sessile or very shortly stalked; pinnules not narrowly oblong, cuneate at basiscopic base, truncate and parallel to costa at acroscopic base; spines of costae and distal part of rachis on adaxial surface prominent   (118)
       
118 (117) Lamina narrowly ovate or lanceolate, acuminate at apex   (119)
+ Lamina ovate or deltoid-ovate, occasionally narrowly ovate, slightly cuspidate or shortly acuminate at apex   (122)
       
119 (118) Costules with prominent spines on adaxial side   (120)
+ Costules without or with very short spines adaxially   (121)
       
120 (119) Rachis and costae shiny, glabrate abaxially; pinnules and lobes usually obtuse at apex, almost not lobed, minutely toothed at margin.   88 A. roseum
+ Rachis and costae densely pubescent abaxially; pinnules and lobes usually rounded-obtuse at apex, shallowly lobed or prominently serrate at margin.   89 A. mengtzeense
       
121 (119) Stipe, rachis, and costae pale purplish red; pinnae shortly stalked (stalk 1-3 mm) or subsessile.   90 A. epirachis
+ Stipe, rachis, and costae stramineous; pinnae sessile.   91 A. pubicostatum
       
122 (118) Rachis and costae glabrate abaxially; pinnae sessile   (123)
+ Rachis and costae pubescent abaxially; pinnae shortly stalked   (124)
       
123 (122) Basal pinnae narrowly lanceolate, acroscopic pinnules subequal to basiscopic pinnules.   92 A. otophorum
+ Basal pinnae ovate-lanceolate, basiscopic pinnules longer than acroscopic pinnules.   93 A. kuratae
       
124 (122) Rachis and costae with short spines on adaxial side, sparsely glandular pubescent abaxially; pinnules separate, short and obtuse at apex.   94 A. arisanense
+ Rachis and costae with long spines adaxially, with dense, pale brown, short glandular hairs abaxially; pinnules dense, acute at apex.   95 A. caudatum

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