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

1. Asplenium Linnaeus, Sp. Pl. 2: 1078. 1753.

铁角蕨属 tie jiao jue shu

Asplenium trichomanes

Credit: Harvard University Herbaria

Acropteris Link; Amesium Newman; Asplenidictyum J. Smith; Asplenium sect. Ceterachopsis J. Smith; A. subg. Thamnopteris C. Presl; Camptosorus Link; Ceterach Willdenow, nom. cons.; Ceterachopsis (J. Smith) Ching; Darea Jussieu; Neottopteris J. Smith; Phyllitis Hill; Schaffneria Fée ex T. Moore; Scolopendrium Adanson; Sinephropteris Mickel; Tarachia C. Presl; Thamnopteris (C. Presl) C. Presl (1851), not Brongniart (1849) [fossil].

Plants epilithic, epiphytic, or terrestrial. Rhizome rarely dorsiventral, creeping, decumbent, or erect, with clathrate scales. Fronds herbaceous to leathery, sometimes subfleshy, remote or (more often) clustered; stipe dull, green to castaneous or black and then often shiny, base terete or semiterete, often becoming sulcate adaxially, rounded to carinate abaxially; lamina simple to 4-pinnate; rachis sulcate adaxially often with raised central supravascular ridge; margin of pinnae or pinnules often decurrent and forming adaxial wings along rachis or costa; acroscopic side of pinnae, pinnules, and ultimate segments often more developed and these then asymmetrical, pseudodimidiate to dimidiate; margin entire to coarsely serrate, each tooth usually with one veinlet, obtuse, mucronate to acute. Veins free, rarely anastomosing, anadromously branching, rarely with a single vein per pinnule or segment, not reaching margin. Sori linear to subelliptic, rarely double, indusiate; indusia thinly membranous to papery, free margin entire to erose, sinuate or fimbriate, rarely with uniseriate glandular hairs; stalks of sporangia long uniseriate, annuli with 20-28 thickened cells. Spores bilateral, elliptic to reniform, monolete, exospore smooth, perispores elaborate and very variable; in sexual plants 64 spores per sporangium. Plants sexual or agamosporous, x = 35, 36.

More than 700 species: subcosmopolitan; 90 species (17 endemic) in China.

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

Uncertain taxa

    Asplenium fangii Ching (Bull. Fan Mem. Inst. Biol., n.s., 1: 276. 1949), described from Sichuan. The type in PE(!) represents a species of Diplazium s.l. Asplenium pekinense Hance var. nanum Christ (Nuovo Giorn. Bot. Ital., n.s., 17(2): 225. 1910), described from Hubei. Type (Silvestri 7, FI) not seen, probably a dwarf form of A. pekinense. Asplenium pseudofalcatum Hillebrand f. obtusatum Rosenstock (Hedwigia 56: 334. 1915), described from Taiwan. Type (Faurie 455bis) not seen, probably a form of A. cuneatiforme. Asplenium pseudofalcatum Hillebrand var. subintegrum Rosenstock (Hedwigia 56: 334. 1915), described from Taiwan. The isotype in MICH (photo!) is probably A. cuneatiforme but is close to A. lobulatum Mettenius ex Kuhn. The relationship of these taxa needs more study. Asplenium wilfordii Mettenius ex Kuhn var. densum Rosenstock (Hedwigia 56: 334. 1915), described from Taiwan. The isotype in NY resembles A. austrochinense, but this taxon should be confirmed for Taiwan. Neottopteris longistipitata R. H. Miao (Acta Sci. Nat. Univ. 1: 99. 1980), described from Hainan.


1 Fronds simple or 2- or 3-forked   (2)
+ Fronds pinnatipartite or more divided   (21)
       
2 (1) Fronds 2- or 3-forked into linear segments   (3)
+ Fronds simple, not forked   (4)
       
3 (2) Average exospore length less than 37 µm; plants sexual diploids: n = 36II or 2n = 72.   1 A. caucasicum
+ Average exospore length more than 37 µm; plants sexual tetraploid: n = 72II or 2n = 144.   2 A. septentrionale
       
4 (2) Veins free, rarely connected near margin, never united into a marginal vein   (5)
+ Veins anastomosing, connected near margin, or united into a marginal vein   (13)
       
5 (4) Sori double (paired).   8 A. komarovii
+ Sori simple   (6)
       
6 (5) Lamina reniform, ca. 1 ?1 cm   (7)
+ Lamina not reniform, larger than ca. 1 ?1 cm   (8)
       
7 (6) Average exospore length 35-40 µm; plants diploid: 2n = 72.   67 A. dolomiticum
+ Average exospore length 41-49 µm; plants tetraploid: 2n = 144.   68 A. ruta-muraria
       
8 (6) Stipe shiny black; lamina less than 10 cm, apex obtuse, margin black.   3 A. speluncae
+ Stipe not shiny black; lamina more than 10 cm, apex acute to acuminate, margin not black   (9)
       
9 (8) Most sori more than 12 mm, at an angle less than 40?to midrib (rachis); lamina margin repand to entire   (10)
+ Most sori less than 12 mm, at an angle of 40?80?to midrib (rachis); lamina margin crenate to sinuate, often notched   (11)
       
10 (9) Fronds lanceolate, 6-10 ?longer than wide, apex acute-acuminate; midrib raised, semiterete adaxially; sori at angle of 30?40?to rachis.   6 A. holosorum
+ Fronds narrowly lanceolate, 10-25 ?longer than wide, apex acuminate-caudate; midrib sulcate adaxially; sori at angle of 15?30?to rachis.   7 A. ensiforme
       
11 (9) Frond stalked; stipe 8-16 cm.   29 A. formosae
+ Frond subsessile; stipe 0.5-6 cm   (12)
       
12 (11) Frond linear to narrowly lanceolate, up to 2 cm wide or more than 13 ?longer than wide.   4 A. scortechinii
+ Frond lanceolate, 2-2.5 cm wide or less than 13 ?longer than wide.   5 A. griffithianum
       
13 (4) Lamina orbicular or ovate, base cordate   (14)
+ Lamina lanceolate to oblanceolate, base cuneate, rarely cordate   (15)
       
14 (13) Rhizome short and erect; fronds clustered; lamina orbicular; most sori double.   9 A. delavayi
+ Rhizome long and creeping; fronds remote; lamina ovate; most sori simple.   see Hymenasplenium cardiophyllum (p. 309)
       
15 (13) Mature fronds less than 20 cm, herbaceous to subleathery, apex usually flagelliform, proliferous and rooting; veins anastomosing, forming 1 or 2 rows of areoles but not ending in a marginal vein.   16 A. ruprechtii
+ Mature fronds more than 20 cm, subleathery to leathery, apex not proliferous; veins ending in a marginal connecting vein (A. subg. Thamnopteris)   (16)
       
16 (15) Frond distinctly spatulate, widest above middle; midrib stramineous to pale brown   (17)
+ Frond lanceolate, widest near middle; midrib pale or grayish brown to brownish black   (18)
       
17 (16) Stipe distinct; midrib semiterete abaxially; lamina reduced at base; plants hexaploid.   10 A. humbertii
+ Stipe very short or absent; midrib keeled abaxially; lamina very gradually reduced into a long broadly winged basal part; plants tetraploid.   11 A. antrophyoides
       
18 (16) Stipe basal scales subulate to very narrowly triangular-ovate, more than 10 ?longer than wide   (19)
+ Stipe basal scales triangular-ovate, less than 10 ?longer than wide   (20)
       
19 (18) Midrib abaxially flat; sori covering up to 1/2 of their subtending vein; perispore winged.   12 A. nidus
+ Midrib abaxially prominent, obtusely carinate; sori covering more than 1/2 of their subtending vein; perispore echinate.   13 A. oblanceolatum
       
20 (18) Stipe basal scales > 15 mm; lamina with many small scales at base; midrib semiterete abaxially.   14 A. antiquum
+ Stipe basal scales < 15 mm; lamina subglabrous at base; midrib obtusely keeled abaxially.   15 A. phyllitidis
       
21 (1) Lamina pinnatipartite-pinnatisect   (22)
+ Lamina 1-4-pinnate   (28)
       
22 (21) Lamina densely covered with scales abaxially; veins anastomosing; indusia reduced to a rim of a few cells.   22 A. ceterach
+ Lamina (sub)glabrous abaxially; veins free; indusia distinct, not reduced to a rim   (23)
       
23 (22) Several pairs of basal pinnae widely separated from more apical ones   (24)
+ Basal pinnae not widely separated from more apical ones (A. subg. Ceterachopsis)   (25)
       
24 (23) Rachis green, without scales.   17 A. castaneoviride
+ Rachis dark brown, with dark filiform scales.   39 A. adnatum
       
25 (23) Segments acute, adaxially often with submarginal gemmae at acroscopic side of pinnae.   21 A. qiujiangense
+ Segments obtuse, without submarginal gemmae at acroscopic side of pinnae   (26)
       
26 (25) Middle segments up to 10 mm wide; mean exospore length < 35 ?.   18 A. dalhousiae
+ Middle segments more than 10 mm wide; mean exospore length > 35 µm   (27)
       
27 (26) Plants up to 20 cm; mean exospore length < 50 ?; plants tetraploid.   19 A. paucivenosum
+ Plants more than 20 cm; mean exospore length > 50 µm; plants octoploid.   20 A. magnificum
       
28 (21) Rhizome long creeping; lamina 1-pinnate   (29)
+ Rhizome erect or shortly creeping; lamina 1-4-pinnate   (30)
       
29 (29) Rhizome less than 0.6 cm in diam., with few scales; venation with several basal basiscopic veins absent.   see Hymenasplenium (p. 308)
+ Rhizome more than 0.6 cm in diam., densely scaly; all basal basiscopic veins present.   37 A. lepturus
       
30 (28) Ultimate pinnules or segments linear, with a single vein; sorus reaching frond margin   (31)
+ Ultimate pinnules or segments not linear, with several veins; sorus rarely reaching frond margin   (37)
       
31 (30) Stipe and rachis shiny dark brown to black on both sides   (32)
+ Stipe and rachis not shiny dark brown to black on both sides   (33)
       
32 (31) Lamina widest at base.   84 A. pulcherrimum
+ Lamina not widest at base.   85 A. cornutissimum
       
33 (31) Plants with minute gemmae on costae or costules.   66 A. tenuifolium
+ Plants not gemmiferous   (34)
       
34 (33) Lamina bipinnate   (35)
+ Lamina 3-pinnatifid to 3-pinnate   (36)
       
35 (34) Rachis apex flagelliform gemmiferous, often rooting and functioning as a runner; pinnae ascending.   86 A. prolongatum
+ Rachis apex not flagelliform; pinnae spreading, attached at right angle to rachis.   87 A. sampsonii
       
36 (34) Pinnae less than 10 cm.   89 A. ritoense
+ Pinnae more than 10 cm.   90 A. trigonopterum
       
37 (30) Lamina 1-pinnate   (38)
+ Lamina pinnate-pinnatifid to 4-pinnate   (61)
       
38 (37) Stipe and rachis shiny, castaneous to dark brown or black   (39)
+ Stipe and rachis dull, not shiny, green, stramineous, or gray-brown   (46)
       
39 (38) Rachis adaxially without brown membranous wings or fingerlike papillae   (40)
+ Rachis adaxially with distinct wings or rows of fingerlike papillae   (41)
       
40 (39) Lamina less than 2 cm wide; rachis semiterete; pinnae 1-2 ?longer than wide.   30 A. kiangsuense
+ Lamina more than 2 cm wide; rachis with deep furrow adaxially; pinnae 2 or more ?longer than wide.   31 A. normale
       
41 (39) Rachis with brown to yellowish brown wings   (42)
+ Rachis adaxially with 2 rows of fingerlike papillae   (44)
       
42 (41) Rachis with 2 adaxial and 1 abaxial wings, often proliferous near apex   24 A. tripteropus
+ Rachis with 2 adaxial wings, abaxial wing absent, never proliferous   (43)
       
43 (42) Plants diploid with average exospore length < 31? and average guard cell length < 39 ?.   25 A. trichomanes
+ Plants tetraploid with average exospore length > 32 µm and average guard cell length > 40 µm.   26 A. quadrivalens
       
44 (41) Rachis gemmiferous at its base; pinnae (slightly) reduced below apex.   27 A. microtum
+ Rachis not gemmiferous; pinnae not reduced near apex   (45)
       
45 (44) Rachis papillae closely set and often coalescent at their base; basal pinnae orbicular-ovate.   28 A. humistratum
+ Rachis papillae separated and rarely coalescent at their base; basal pinnae hastate-deltoid.   29 A. glanduliserrulatum
       
46 (38) Costa raised and semiterete on adaxial surface of pinna   (47)
+ Costa flat or depressed (grooved) on adaxial surface of pinna   (50)
       
47 (46) Frond apex a single pinna, conform with lateral ones.   32 A. formosae
+ Frond apex a pinnatisect or pinnatifid terminal segment different from lateral pinnae   (48)
       
48 (47) Frond apex of 1 or 2 basal segments attached to single terminal pinna; indusia not rolling back at maturity; costa with first basiscopic vein lacking and more distal veins simple.   33 A. matsumurae
+ Frondapex pinnatifid and consisting of several confluent segments; indusia rolling back at maturity; costa with all veins present and forking   (49)
       
49 (48) Pinnae obtuse, usually less than 5 cm.   88 A. tenerum
+ Pinnae acuminate to caudate, usually larger.   34 A. wrightii
       
50 (46) Frond apex similar to one lateral pinna   (51)
+ Frond apex pinnatifid and not similar to a lateral pinna   (54)
       
51 (50) Plants less than 10 cm tall   (52)
+ Plants more than 10 cm tall   (53)
       
52 (51) Average exospore length 35-40 µm; plants diploid: 2n = 72.   67 A. dolomiticum
+ Average exospore length 41-49 µm; plants tetraploid: 2n = 144.   68 A. ruta-muraria
       
53 (51) Stipe and rachis pale yellow-green to gray; veins irregularly anastomosing near margin.   35 A. finlaysonianum
+ Stipe and rachis brown to dark brown; veins free.   36 A. polyodon
       
54 (50) Lamina more than 10 cm wide   (55)
+ Lamina less than 10 cm wide   (56)
       
55 (54) Frond herbaceous; rhizome creeping; rachis often gemmiferous near apex; spores pale, perispore echinate.   38 A. trapezoideum
+ Frond papery; rhizome erect; rachis not gemmiferous; spores dark brown-black, perispore lophate.   47 A. saxicola
       
56 (54) Rachis dark brown to black, with many dark brown to blackish scales with long filiform apex   (57)
+ Rachis gray-stramineous to green, with few narrowly triangular, reddish brown scales without filiform apex   (58)
       
57 (56) Frond shorter than 30 cm, less than 3 cm wide; pinnae oblong, obtuse.   40 A. asterolepis
+ Fronds longer than 30 cm, more than 3 cm wide; pinnae narrowly ovate, acute   41 A. crinicaule
       
58 (56) Lamina less than 1.5 cm wide; stipe and rachis green, never gemmiferous.   23 A. viride
+ Lamina more than 1.5 cm wide; stipe dull grayish brown, rachis green in apical part and occasionally gemmiferous   (59)
       
59 (58) Gemmae on adaxial surface near pinna apex; scales costate with dark brown central zone and pale hyaline margin; perispore cristate-alate (with narrow crests on thin wings).   44 A. gueinzianum
+ Gemmae on rachis or close to rachis on pinna stalk; scales not costate; perispore costate (with rounded crest on broad ridges)   (60)
       
60 (59) Plants tetraploid: 2n = 144; average exospore length less than 36µm.   42 A. indicum
+ Plants octoploid: 2n = 288; average exospore length more than 36µm.   43 A. yoshinagae
       
61 (37) Stipe entirely shiny dark brown to black.   83 A. coenobiale
+ Stipe not entirely shiny dark brown to black   (62)
       
62 (61) Stipe dark gray to dark brown   (63)
+ Stipe green, abaxially at base and higher up often castaneous to black   (82)
       
63 (62) Fronds pinnate-pinnatifid to almost 2-pinnate   (64)
+ Fronds more divided, up to 4-pinnatifid   (70)
       
64 (63) Gemmae present   (65)
+ Gemmae absent   (66)
       
65 (64) Bud on adaxial surface of pinnae.   44 A. gueinzianum
+ Bud on adaxial surface of rachis below apex.   50 A. cuneatiforme
       
66 (64) Rachis densely scaly   (67)
+ Rachis not densely scaly   (68)
       
67 (66) Fronds widest near base; stipe more than 5 cm; scales with filiform apical tail.   45 A. aethiopicum
+ Fronds widest near middle; stipe less than 5 cm; scales without filiform apical tail.   51 A. rockii
       
68 (66) Plants less than 20 cm tall, usually less than 10 pinnae pairs.   48 A. oldhamii
+ Plants more than 20 cm tall, usually more than 10 pinnae pairs   (69)
       
69 (68) Fronds herbaceous; pinna stalk less than 4 mm.   46 A. pseudopraemorsum
+ Fronds leathery; pinna stalk more than 4 mm.   47 A. saxicola
       
70 (63) Fronds 3-pinnate or 4-pinnatifid   (71)
+ Fronds 2-pinnate to 3-pinnatifid   (75)
       
71 (70) Plants less than 50 cm   (72)
+ Plants more than 50 cm   (73)
       
72 (71) Ultimate soriferous segment cuneiform, with several veins and sori.   56 A. wilfordii
+ Ultimate soriferous segment linear with 1 vein and 1 sorus.   89 A. ritoense
       
73 (71) Fronds herbaceous; indusia elliptic, ca. 4 mm; indusium margin with glandular hairs.   55 A. bullatum
+ Fronds papery to subleathery; indusia linear, most than 4 mm; indusium margin without glandular hairs   (74)
       
74 (73) Pinnules broadly ovate, less than 2 ?longer than wide.   57 A. sublaserpitiifolium
+ Pinnules triangular-ovate, more than 2 ?longer than wide.   58 A. pseudolaserpitiifolium
       
75 (70) Gemmae present on rachis   (76)
+ Gemmae absent   (77)
       
76 (75) Pinnae with more than 3 pairs of free pinnules.   49 A. affine
+ Pinnae with less than 3 pairs of free pinnules.   50 A. cuneatiforme
       
77 (75) Stipe less than 5 cm; rachis densely scaly.   51 A. rockii
+ Stipe more than 5 cm; rachis not densely scaly   (78)
       
78 (77) Largest pinnae with less than 3 pairs of free pinnules, most decurrent on costa   (79)
+ Largest pinnae with more than 3 pairs of free pinnules   (80)
       
79 (78) Sori more than 5 mm.   47 A. saxicola
+ Sori less than 5 mm.   52 A. austrochinense
       
80 (78) Plants less than 50 cm tall.   53 A. hainanense
+ Plants more than 50 cm tall   (81)
       
81 (80) More than 5 free pinnules per pinna; veins close together and almost parallel; free margin of indusium glabrous.   54 A. subspathulinum
+ Less than 5 free pinnules per pinna; venation different; free margin of indusium with glandular hairs.   55 A. bullatum
       
82 (62) Fronds lanceolate, reduced at base   (83)
+ Fronds ovate-triangular, not reduced or not much reduced at base   (87)
       
83 (82) Pinnae with minute gemma or plantlet at apex   (84)
+ Pinnae not gemmiferous at apex   (85)
       
84 (83) Average exospore length less than 29 µm; plants diploid: 2n = 72.   59 A. lushanense
+ Average exospore length more than 29 µm; plants tetraploid: 2n = 144   60 A. exiguum
       
85 (83) Fronds thinly herbaceous; stipe and rachis with thin, green lateral wings   61 A. incisum
+ Fronds firmly herbaceous; stipe and rachis without thin, green wings   (86)
       
86 (85) Marginal teeth usually slender, obtuse to subacute.   62 A. nesii
+ Marginal teeth short, subacute to mucronate.  

A. fontanum

       
87 (82) Small gemmae present on pinnae stalk, costa, costule, or rachis tip   (88)
+ Gemmae absent   (90)
       
88 (87) Pinnae usually with more than 3 pairs of free pinnules; gemmae on costa or costule.   66 A. tenuifolium
+ Pinnae with less than 3 pairs of free pinnules; gemmae on pinnae stalk or on flagelliform apex   (89)
       
89 (88) Lamina triangular, without extended gemmiferous rachis.   64 A. capillipes
+ Lamina oblong-linear with flagelliform gemmiferous apex.   65 A. fugax
       
90 (87) Fronds deltoid to broadly triangular   (91)
+ Fronds triangular to ovate; indusial margin not fimbriate   (93)
       
91 (90) Margin of indusium not fimbriate.   69 A. interjectum
+ Margin of indusium fimbriate (with long uniseriate hairs)   (92)
       
92 (91) Average exospore length 35-40 µm; plants diploid: 2n = 72.   67 A. dolomiticum
+ Average exospore length 41-49 µm; plants tetraploid: 2n = 144.   68 A. ruta-muraria
       
93 (90) Stipe green or only brown at very base; rachis entirely green   (94)
+ Stipe and often base of rachis brown on abaxial side   (95)
       
94 (93) Fronds thinly herbaceous; average exospore length less than 32 µm; plants diploid: 2n = 72.   81 A. sarelii
+ Fronds firmly herbaceous to subleathery; average exospore length more than 32 µm; plants tetraploid: 2n = 144.   82 A. pekinense
       
95 (93) Fronds 3-pinnate at base; pinnae with more than one free acroscopic pinnule   (96)
+ Fronds 2-pinnate at base; pinnae with 1 free acroscopic pinnule   (99)
       
96 (95) Plants diploid; average exospore length less than 32 ?.   70 A. tenuicaule
+ Plants polyploid; average exospore length more than 32 ?   (97)
       
97 (96) Lamina triangular; stipe base swollen.   79 A. adiantum-nigrum
+ Lamina narrowly triangular to ovate; stipe base not swollen   (98)
       
98 (97) Pinna stalk wiry; perispore with many wide pori (reticulate) and almost no ridges.   76 A. kansuense
+ Pinna stalk not wiry; perispore with ridges and few minute pori (cristate).   80 A. anogrammoides
       
99 (95) Plants diploid; average exospore length less than 32 ?   (100)
+ Plants polyploid; average exospore length more than 32 ?   (101)
       
100 (99) Pinna stalk wiry; marginal teeth broad, mucronate; average exospore length 29-32 ?; perispore reticulate or cristate.   70 A. tenuicaule
+ Pinna stalk not wiry; marginal teeth slender, acute; average exospore length 24-27 ?; perispore alate.   71 A. semivarians
       
101 (99) Lamina triangular, widest at base; largest pinnae broadly ovate-deltoid   (102)
+ Lamina ovate, widest just above base; largest pinnae ovate to narrowly triangular   (103)
       
102 (101) Plants hexaploid; average exospore length 38-42 ?.   72 A. aitchisonii
+ Plants tetraploid; average exospore length 33-36 ?.   73 A. neovarians
       
103 (101) Lamina thin, membranous; largest pinnae triangular to narrowly triangular.   74 A. mae
+ Lamina more firm, herbaceous; largest pinnae ovate to triangular   (104)
       
104 (103) Lamina acuminate to shortly caudate; scales with inner part of cell walls warty.   75 A. kukkonenii
+ Lamina not shortly caudate; scales with inner part of cell walls smooth   (105)
       
105 (104) Lamina firmly herbaceous to subleathery; basal pinnae deltoid to flabellate.   78 A. altajense
+ Lamina herbaceous; basal pinnae oval-triangular   (106)
       
106 (105) Pinna stalk wiry; perispore with many wide pori (reticulate) and almost no ridges.   76 A. kansuense
+ Pinna stalk not wiry; perispore with ridges and few minute pori (cristate).   77 A. varians

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