铁角蕨科 tie jiao jue ke
Lin Youxing (林尤兴)[1]; Ronald Viane[2]
Ferns middle sized to small, predominantly lithophytic or terrestrial, but also many epiphytic. Rhizome creeping, decumbent, or erect, radial or dorsiventral (Hymenasplenium), sometimes massive and bearing detritus-collecting fronds, scaly; scales narrowly triangular, clathrate with anticlinal walls thickened and periclinal walls translucent, often costate and with a central zone of cells with thicker anticlinal walls, in some taxa becoming completely opaque, rarely whole scale black and opaque, margins entire, dentate, or with short or long outgrowths (fimbriate scales), often with a terminal glandular cell, both surfaces glabrous. Fronds remote (in China mainly in Hymenasplenium) or clustered, spirally arranged or in two abaxial rows (Hymenasplenium), herbaceous or leathery to subfleshy, with scales similar to those on rhizome or gradually reduced in size (mini-scales) and filiform to hastate-stellate, usually with small, 3- or 4-celled uniseriate hairs terminating in a gland, rarely glandular or completely glabrous, occasionally gemmiferous with buds often on, but not restricted to, rachis or at pinna apex; stipe usually distinct, not articulate, rarely forming a trophopod (Asplenium adiantum-nigrum) or persistent (Hymenasplenium), distinctly scaly at base, green to grayish brown, in some species castaneous to black and then often shiny, rounded to grooved (sulcate) adaxially, with two vascular bundles at its base, upward near rachis combined into one X-shaped bundle. Lamina outline variable, from simple to 4-pinnate, apical pinnae usually reduced and confluent into a pinnatifid apex, rarely conforming to lateral pinnae (e.g., A. formosae), usually branching anadromously; rachis structure variable, adaxially often sulcate, with or without central supravascular ridge, with or without lateral wings derived from decurrent frond margins, rarely with an abaxial wing (A. tripteropus); pinnae and other divisions often more strongly developed in direction of apex, ultimate pinnules or segments often rhombic, trapeziform, or cuneiform with asymmetric base, rarely dimidiate, margin rarely entire, more often repand or sinuate to crenate or serrate, each tooth usually with a single vein. Veins mostly free, anadromously branching, veinlets usually not reaching margin, with or without terminal hydathode, rarely anastomosing near margin or with a marginal connecting vein (A. subg. Thamnopteris C. Presl); some species with a dissected lamina have one single vein per ultimate segment. Sori single, rarely double on two different veins (A. subg. Phyllitis (Hill) Jermy & Viane), or J-shaped (e.g., A. fontanum), linear to subelliptic, attached along one side of a veinlet, usually indusiate (indusium reduced in A. subg. Ceterach (Willdenow) Vida ex Bir, Fraser-Jenkins & Lovis); indusia thinly membranous to thick and firm, usually flat but occasionally rolling back at maturity, free margin entire to erose, sinuate, fimbriate, glandular, or rarely with mini-scales, attached laterally along a veinlet, opening toward costa, costule, or vein from which soriferous vein originated, in some species with a dissected lamina seemingly opening toward margin (A. prolongatum). Sporangia with a vertical annulus of ca. 20 thickened cells, stalk uniseriate, not glandular. Spores usually 64 per sporangium, bilateral, elliptic to reniform, monolete; exospore smooth, perispores elaborate and very variable, consisting of three layers, outer with ridges (lophate), spines (echinate), or with large pori (reticulate) and no ridges or spines, or a combination. Plants sexual or agamosporous and then often with only 32 spores per sporangium, x = (35) 36 in Asplenium, but (36, 38)39 in Hymenasplenium.
Two genera and ca. 700 species: subcosmopolitan, but mainly in montane and (sub)tropical regions; two genera and 108 species and species complexes (Hymenasplenium) (26 endemic) in China.
Wu Shiew-hung. 1999. Aspleniaceae. In: Wu Shiew-hung, ed., Fl. Reipubl. Popularis Sin. 4(2): 1153.
1a. Rhizome erect or shortly creeping, if long creeping then usually more than 0.6 cm in diam. and densely scaly throughout; fronds remote or clustered, lamina simple to 4-pinnate; costa usually with all basal basiscopic veins present ..................................................................................................... 1. Asplenium
1b. Rhizome long creeping,
usually less than 0.6 cm in diam. and with very few scales except near apex,
fronds remote, lamina rarely simple, usually 1-pinnate; costa usually with
several (rarely one) basal basiscopic veins absent
2.
Hymenasplenium
铁角蕨属 tie jiao jue shu
Plants lithophytic, 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 2028 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: [worldwide distribution?]; 90 species (18 endemic) in China.
1a. Frond simple or 2 or 3-forked.
2a. Fronds 2- or 3-forked into linear segments.
3a. Average exospore length less than 37 ΅m; plants sexual diploids: n = 36II or 2n = 72 1. A. caucasicum
3b. Average exospore length more than 37 ΅m; plants sexual tetraploid: n = 72II or 2n = 144 2. A. septentrionale
2b. Fronds simple, not forked.
4a. Veins free, rarely connected near margin, never united into a marginal vein.
5a. Sori double (paired) ............................................................................... 8. A. komarovii
5b. Sori simple.
6a. Lamina reniform, ca. 1 Χ 1 cm.
7a. Average exospore length 3540 ΅m; plant diploid: 2n = 72 .......... 67. A. dolomiticum
7b. Average exospore length 4149 ΅m; plant tetraploid: 2n = 144
................................................................................................... 68.
A. ruta-muraria
6b. Lamina not reniform, larger than ca. 1 Χ 1 cm.
8a. Stipe shiny black; lamina less than 10 cm, apex obtuse,
margin black
.......................................................................................................... 3.
A. speluncae
8b. Stipe not shiny black; lamina more than 10 cm, apex acute to acuminate, margin not black.
9a. Most sori more than 12 mm, at an angle less than 40° to midrib (rachis); lamina margin repand to entire.
10a. Fronds lanceolate, 610 Χ longer than wide, apex
acute-acuminate; midrib raised, semiterete adaxially; sori at angle of 3040°
to rachis
6.
A. holosorum
10b. Fronds narrowly lanceolate, 1025 Χ longer than wide, apex acuminate-caudate; midrib sulcate adaxially; sori at angle of 1530° to rachis ..................................... 7. A. ensiforme
9b. Most sori less than 12 mm, at an angle of 4080° to midrib (rachis); lamina margin crenate (notched) to sinuate.
11a. Frond stalked; stipe 816 cm ..................................... 29. A. loriceum[formosae?]
11b. Frond subsessile; stipe 0.56 cm.
12a. Frond linear to narrowly lanceolate, up to 2 cm wide or more than 13 Χ longer than wide 4. A. scortechinii
12b. Frond lanceolate, 22.5 cm wide or less than 13 Χ longer
than wide
.............................................................................................. 5.
A. griffithianum
4b. Veins anastomosing, connected near margin, or united into a marginal vein.
13a. Lamina orbicular or ovate, base cordate.
14a. Rhizome short and erect; fronds clustered; lamina orbicular; most sori double 9. A. delavayi
14b. Rhizome long and creeping; fronds remote; lamina ovate;
most sori simple
see Hymenasplenium cardiophyllum (p. xxx)
13b. Lamina lanceolate to oblanceolate, base cuneate, rarely cordate.
15a. Mature fronds less than 20 cm; herbaceous to subleathery; apex usually flagelliform, proliferous and rooting; veins anastomosing, forming 1 or 2 rows of areolae but not ending in a marginal vein 16. A. ruprechtii
15b. Mature fronds more than 20 cm, subleathery to leathery, apex not proliferous; veins ending in a marginal connecting vein (A. subg. Thamnopteris).
16a. Frond distinctly spatulate, widest above middle; midrib stramineous to pale brown.
17a. Stipe distinct; midrib semiterete abaxially; lamina reduced at base; plants hexaploid 10. A. humbertii
17b. Stipe very short or absent; midrib keeled abaxially; lamina very gradually reduced into a long broadly winged basal part; plants tetraploid ............................. 11. A. antrophyoides
16b. Frond lanceolate, widest near middle; midrib pale or grayish brown to brownish black.
18a. Stipe basal scales subulate to very narrowly triangular-ovate, more than 10 Χ longer than wide.
19a. Midrib abaxially flat to slightly semiterete; sori covering up to 1/2 of their subtending vein; perispore winged ................................................................................ 12. A. nidus
19b. Midrib abaxially prominent obtusely carinate; sori covering more than 1/2 of their subtending vein; perispore echinate ....................................................... 13. A. oblanceolatum
18b. Stipe basal scales triangular-ovate, less than 10 Χ longer than wide.
20a. Stipe basal scales > 15 mm; lamina with many small scales at base; midrib semiterete abaxially 14. A. antiquum
20b. Stipe basal scales < 15 mm; lamina subglabrous at base; midrib obtusely keeled abaxially 15 A. pyllitidis
1b. Fronds pinnatipartite or more divided.
21a. Lamina pinnatipartite-pinnatisect.
22a. Lamina densely covered with scales abaxially; veins anastomosing; indusia reduced to a rim of a few cells .......................................................................................................... 22. A. ceterach
22b. Lamina (sub)glabrous abaxially; veins free; indusia distinct, not reduced to a rim.
23a. Several pairs of basal pinnae widely separated from more apical ones.
24a. Rachis green, without scales ...................................................... 17. A. castaneo-viride
24b. Rachis dark brown, with dark filiform scales ......................................... 39. A. adnatum
23b. Basal pinnae not widely separated from more apical ones (A. subg. Ceterachopsis).
25a. Segments acute, adaxially often with submarginal gemmae at acroscopic side of pinnae 21. A. qiujiangense
25b. Segments obtuse, without submarginal gemmae at acroscopic side of pinnae.
26a. Middle segments up to 10 mm wide; mean exospore length
< 35 ΅m
....................................................................................................... 18.
A. dalhousiae
26b. Middle segments more than 10 mm wide; mean exospore length > 35 ΅m.
27a. Plants up to 20 cm; mean exospore length < 50 ΅m;
plants tetraploid
............................................................................................... 19.
A. paucivenosum
27b. Plants more than 20 cm; mean exospore length > 50 ΅m; plants octoploid 20. A. magnificum
21b. Lamina 14-pinnate.
28a. Rhizome long creeping; lamina 1-pinnate.
29a. Rhizome less than 0.6 cm in diam., with few scales; venation with several basal basiscopic veins absent ......................................................................................... see Hymenasplenium (p. xxx)
29b. Rhizome more than 0.6 cm in diam., densely scaly; all basal basiscopic veins present 37. A. lepturus
28b. Rhizome erect or shortly creeping; lamina 14-pinnate.
30a. Ultimate pinnules or segments linear, with a single vein; sorus reaching frond margin.
31a. Stipe and rachis shiny dark brown to black on both sides.
32a. Lamina widest at base ............................................................... 84. A. pulcherrimum
32b. Lamina not widest at base ........................................................ 85. A. cornutissimum
31b. Stipe and rachis not shiny dark brown to black on both sides.
33a. Plants with minute gemmae on costae or costules ........................... 66. A. tenuifolium
33b. Plants not gemmiferous.
34a. Lamina bipinnate.
35a. Apex flagelliform gemmiferous, often rooting and functioning as a runner; pinnae ascending 86. A. prolongatum
35b. Apex not flagelliform; pinnae spreading, attached at right angle to rachis 87. A. sampsonii
34b. Lamina 3-pinnatifid to 3-pinnate.
36a. Pinnae less than 10 cm ................................................................... 89. A. ritoense
36b. Pinnae more than 10 cm ....................................................... 90. A. trigonopterum
30b. Ultimate pinnules or segments not linear, with several veins; sorus rarely reaching frond margin.
37a. Lamina 1-pinnate.
38a. Stipe and rachis shiny, castaneous to dark brown or black.
39a. Rachis adaxially without brown membranous wings or fingerlike papillae.
40a. Lamina less than 2 cm wide; rachis semiterete; pinnae 12 Χ longer than wide 30. A. kiangsuense
40b. Lamina more than 2 cm wide; rachis with deep furrow adaxially; pinnae 2 or more times longer than wide ....................................................................................... 31. A. normale
39b. Rachis adaxially with distinct wings or lateral rows of fingerlike papillae.
41a. Rachis with brown to yellowish brown wings.
42a. Rachis with two adaxial and one abaxial wings, often proliferous near apex 24. A. tripteropus
42b. Rachis with two adaxial wings, abaxial wing absent, never proliferous.
43a. Plants diploid with average exospore length < 31΅m and average guard cell length < 39 ΅m 25. A. trichomanes
43b. Plants tetraploid with average exospore length >32 ΅m and average guard cell length > 40 ΅m 26. A. lovisii
41b. Rachis adaxially with lateral row of fingerlike papillae.
44a. Rachis gemmiferous at its base; pinnae (slightly)
reduced below apex
................................................................................................. 27.
A. microtum
44b. Rachis not gemmiferous; pinnae not reduced near apex.
45a. Rachis papillae closely set and often coalescent at their base; basal pinnae orbicular-ovate 28. A. humistratum
45b. Rachis papillae separated and rarely coalescent at their base; basal pinnae hastate-deltoid 29. A. glanduliserrulatum
38b. Stipe and rachis dull, not shiny, green, stramineous, or gray-brown.
46a. Costa raised and semiterete on adaxial surface of pinna.
47a. Frond apex a single pinna, conforming to lateral ones .................... 32. A. formosae
47b. Frond apex a pinnatisect or pinnatifid terminal segment different from lateral pinnae.
48a. 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
48b. Apex pinnatifid and consisting of several confluent segments; indusia rolling back at maturity; costa with all veins present and forking.
49a. Pinnae obtuse, usually less than 5 cm ......................................... 88. A. tenerum
49b. Pinnae acuminate to caudate, pinnae larger ................................. 34. A. wrightii
46b. Costa flat or depressed (grooved) on adaxial surface of pinna.
50a. Frond apex similar to one lateral pinna.
51a. Plants less than 10 cm tall.
52a. Average exospore length 3540 ΅m; plant diploid: 2n = 72
67.
A. dolomiticum
52b. Average exospore length 4149 ΅m; plant tetraploid: 2n = 144
68.
A. ruta-muraria
51b. Plants more than 10 cm tall.
53a. Stipe and rachis pale yellow-green to gray; veins irregularly anastomosing near margin 35. A. finlaysonianum
53b. Stipe and rachis brown to dark brown; veins free ..................... 36. A. polyodon
50b. Frond apex pinnatifid and not similar to a lateral pinna.
54a. Lamina more than 10 cm wide.
55a. Frond herbaceous; rhizome creeping; rachis often gemmiferous near apex; spores pale, perispore echinate ............................................................ 38. A. trapezoideum
55b. Frond papery; rhizome erect; rachis not gemmiferous; spores dark brown-black, perispore lophate ..................................................................................... 47. A. saxicola
54b. Lamina less than 10 cm wide.
56a. Rachis dark brown to black, with many dark brown to blackish scales with long filiform apex.
57a. Frond shorter than 30 cm, less than 3 cm wide; pinnae oblong, obtuse 40. A. asterolepis
57b. Fronds longer than 30 cm, more than 3 cm wide; pinnae narrowly ovate, acute 41. A. crinicaule
56b. Rachis gray-stramineous to green, with few narrowly triangular, reddish brown scales without filiform apex.
58a. Lamina less than 1.5 cm wide; stipe and rachis green, never gemmiferous 23. A. viride
58b. Lamina more than 1.5 cm wide; stipe dull grayish brown, rachis green in apical part and occasionally gemmiferous.
59a. 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
59b. Gemmae on rachis or close to rachis on pinna stalk; scales not costate; perispore costate (with rounded crest on broad ridges).
60a. Plants tetraploid: 2n = 144; average exospore length less than 36΅m 42. A. indicum
60b. Plants octoploid: 2n = 288; average exospore length more than 36΅m 43. A. yoshinagae
37b. Lamina pinnate-pinnatifid to 4-pinnate.
61a. Stipe entirely shiny dark brown to black .......................................... 83. A. coenobiale
61b. Stipe not entirely shiny dark brown to black.
62a. Stipe dark gray to dark brown.
63a. Fronds pinnate-pinnatifid to almost 2-pinnate.
64a. Gemmae present.
65a. Bud on adaxial surface of pinnae ........................................ 44. A. gueinzianum
65b. Bud on adaxial surface of rachis below apex ...................... 50. A. cuneatiforme
64b. Gemmae absent.
66a. Rachis densely scaly.
67a. Fronds widest near base; stipe more than 5 cm; scales with filiform apical tail 45. A. aethiopicum
67b. Fronds widest near middle; stipe less than 5 cm; scales without filiform apical tail 51. A. rockii
66b. Rachis not densely scaly.
68a. Plants less than 20 cm tall; usually less than 10
pinnae pairs
.............................................................................................. 48.
A. oldhamii
68b. Plants more than 20 cm tall, usually more than 10 pinnae pairs.
69a. Fronds herbaceous; pinna stalk less than 4 mm ... 46. A. pseudopraemorsum
69b. Fronds leathery; pinna stalk more than 4 mm ......................... 47. A. saxicola
63b. Fronds more divided, up to 4-pinnatifid.
70a. Fronds 3-pinnate or 4-pinnatifid.
71a. Plants less than 50 cm.
72a. Ultimate soriferous segments cuneiform, with several veins and sori 56. A. wilfordii
72b. Ultimate soriferous segment linear with 1 vein and 1 sorus ........ 89. A. ritoense
71b. Plants more than 50 cm.
73a. Fronds herbaceous; indusium elliptic, ca. 4 mm; indusium margin with glandular hairs 55. A. bullatum
73b. Fronds papery to subleathery; indusia linear, most than 4 mm; indusium margin without glandular hairs.
74a. Pinnules broadly ovate, less than twice longer as wide
57.
A. sublaserpitiifolium
74b. Pinnules triangular-ovate, more than twice longer as
wide
58. A. pseudolaserpitifolium
70b. Fronds 2-pinnate to 3-pinnatifid.
75a. Gemmae present on rachis.
76a. Pinnae with more than 3 pairs of free pinnules .............................. 49. A. affine
76b. Pinnae with less than 3 pairs of free pinnules .................... 50. A. cuneatiforme
75b. Gemmae absent.
77a. Stipe less than 5 cm; rachis densely scaly .................................... 51. A. rockii
77b. Stipe more than 5 cm; rachis not densely scaly.
78a. Largest pinnae with less than 3 pairs of free pinnules, most decurrent on costa.
79a. Sori more than 5 mm ......................................................... 47. A. saxicola
79b. Sori less than 5 mm ................................................. 52. A. austrochinense
78b. Largest pinnae with more than 3 pairs of free pinnules.
80a. Plants less than 50 cm tall .............................................. 53. A. hainanense
80b. Plants more than 50 cm tall.
81a. More than 5 free pinnules per pinna; veins close together and almost parallel; free margin of indusium glabrous ...................................................... 54. A. yui
81b. Less than 5 free pinnules per pinna; venation different; free margin of indusium with glandular hairs ................................................................ 55. A. bullatum
62b. Stipe green, abaxially at base and higher up often castaneous to black.
82a. Fronds lanceolate, reduced at base.
83a. Pinnae with minute gemma or plantlet at apex.
84a. Average exospore length less than 29 ΅m; diploid: 2n = 72
59.
A. lushanense
84b. Average exospore length more than 29 ΅m; tetraploid: 2n = 144
60.
A. exiguum
83b. Pinnae not gemmiferous at apex.
85a. Fronds thinly herbaceous; stipe and rachis with thin, green lateral wings 61. A. incisum
85b. Fronds firmly herbaceous; stipe and rachis without thin, green wings.
86a. Marginal teeth usually slender, obtuse to subacute ......................... 62. A. nesii
86b. Marginal teeth short, subacute to mucronate .......................... 63. A. fontanum
82b. Fronds ovate-triangular, not reduced or not much reduced at base.
87a. Small gemmae present on pinnae stalk, costa, costule, or rachis tip.
88a. Pinnae usually with more than 3 pairs of free pinnules; gemmae on costa or costule 66. A. tenuifolium
88b. Pinnae with less than 3 pairs of free pinnules; gemmae on pinnae stalk or on flagelliform apex.
89a. Lamina triangular, without extended gemmiferous rachis ......... 64. A. capillipes
89b. Lamina oblong-linear with flagelliform gemmiferous apex ............. 65. A. fugax
87b. Gemmae absent.
90a. Fronds deltoid to broadly triangular.
91a. Margin of indusium not fimbriate ........................................ 69. A. interjectum
91b. Margin of indusium fimbriate (with long uniseriate hairs).
92a. Average exospore length 3540 ΅m; plant diploid: 2n = 72
67.
A. dolomiticum
92b. Average exospore length 4149 ΅m; plant tetraploid: 2n = 144
68.
A. ruta-muraria
90b. Fronds triangular to ovate; indusial margin not fimbriate.
93a. Stipe green or only brown at very base; rachis entirely green.
94a. Fronds thinly herbaceous; average exospore length less than 32 ΅m; diploid: 2n = 72 81. A. sarrelii
94b. Fronds firmly herbaceous to subleathery; average exospore length more than 32 ΅m; tetraploid: 2n = 144 ........................................................... 82. A. pekinense
93b. Stipe and often base of rachis brown on abaxial side.
95a. Fronds 3-pinnate at base; pinnae with more than one free acroscopic pinnule.
96a. Plants diploid; average exospore length less than 32 ΅m
....................................................................................... 70.
A. tenuicaule
96b. Plants polyploid; average exospore length more than 32 ΅m.
97a. Lamina triangular; stipe base swollen ................. 79. A. adiantum-nigrum
97b. Lamina narrowly triangular to ovate; stipe base not swollen.
98a. Pinna stalk wiry; perispore with many wide pori (reticulate) and almost no ridges 76. A. kansuense
98b. Pinna stalk not wiry; perispore with ridges and few minute pori (cristate) 80. A. anogrammoides
95b. Fronds 2-pinnate at base; pinnae with 1 free acroscopic pinnule.
99a. Plants diploid; average exospore length less than 32 ΅m.
100a. Pinna stalk wiry; marginal teeth broad, mucronate;
average exospore length 2932 ΅m; perispore reticulate or cristate
70.
A. tenuicaule
100b. Pinna stalk not wiry; marginal teeth slender, acute; average exospore length 2427 ΅m; perispore alate ........................................................... 71. A. semivarians
99b. Plants polyploid; average exospore length more than 32 ΅m.
101a. Lamina triangular, widest at base; largest pinnae broadly ovate-deltoid.
102a. Plants hexaploid; average exospore length 3842 ΅m
.................................................................................. 72.
A. aitchisonii
102b. Plants tetraploid; average exospore length 3336 ΅m
................................................................................. 73.
A. neovarians
101b. Lamina ovate, widest just above base; largest pinnae ovate to narrowly triangular.
103a. Lamina thin, membranous; largest pinnae triangular to narrowly triangular 74. A. mae
103b. Lamina more firm, herbaceous; largest pinnae ovate to triangular.
104a. Lamina acuminate to shortly caudate; scales with inner part of cell walls warty 75. A. kukkonenii
104b. Lamina not shortly caudate; scales with inner part of cell walls smooth.
105a. Lamina firmly herbaceous to subleathery; basal pinnae deltoid to flabellate 78. A. altajense
105b. Lamina herbaceous; basal pinnae oval-triangular.
106a. Pinna stalk wiry; perispore with many wide pori (reticulate) and almost no ridges ........................................................................... 76. A. kansuense
106b. Pinna stalk not wiry; perispore with ridges and few minute pori (cristate) 77. A. varians
Chinese pinyin
Asplenium septentrionale (Linnaeus) Hoffmann subsp. caucasicum Fraser-Jenkins & Lovis, Notes Roy. Bot. Gard. Edinburgh 38: 281. 1980.
Plants 815 cm tall. Rhizome shortly creeping to ascending, apex scaly; scales dark brown, narrowly triangular to subulate, entire to denticulate, at base fimbriate. Fronds caespitose; stipe shiny castaneous only at base, abruptly green toward rachis, 610 cm, 23 Χ as long as lamina, with unicellular glands or subglabrous, apex 2- or 3-forked; segments linear, 23 Χ 0.10.15 cm, base gradually decurrent onto stipe, minutely forked again at apex. Fronds without distinct rachis, green, other veins slender and subparallel to central axis, one veinlet per ultimate segment. Fronds herbaceous-leathery, grass-green. Sori 15 per segment, linear, 12 cm, on acroscopic veins and close to costa, at maturity covering entire surface; indusia brown, linear, membranous, free margin often with unicellular glands, entire-sinuate, opening toward main vein (costa), persistent but reflexed and covered by sporangia at maturity. Spores with lophate perispore, average exospore length 3236 ΅m. Plants sexual diploid: 2n = 72.
In crevices of non-calcareous (often granitic) rocks in open or partial shade; below 4100 m. ?Taiwan, Xinjiang, Xizang [Pakistan, Russia; SW Asia (Abkhazia/Georgia, Azerbaijan, Iran, Turkey), E Europe].
The type specimen is from Abkhazia/Georgia.
Asplenium caucasicum is the diploid ancestor of the more common autotetraploid A. septentrionale (see below). This taxon is best known from the Caucasus region, but microcharacters have shown its presence in Xinjiang. A recent chromosome count from Xizang confirms its presence in China. A doubling of its genome has led to the origin of autotetraploid A. septentrionale. In places where both taxa grow together, their triploid sterile hybrid, A. Χdirense Viane & Reichstein, can be expected.
Asplenium sasakii was distinguished from A. septentrionale by its narrower fronds (Tagawa, [??Acta Phytotax. Geobot. 10: ??]. 1941); it may be an earlier name for A. caucasicum if Taiwanese plants turn out to be diploid. However, at present, no chromosome counts or flow-cytometric data are available for Taiwanese plants.
叉叶铁角蕨 cha ye tie jiao jue
Acrostichum septentrionale Linnaeus, Sp. Pl. 2: 1068. 1753; Amesium sasakii Hayata; Asplenium sasakii (Hayata) Makino & Nemoto; A. septentrionale var. sasakii (Hayata) C. Christensen.
Plants 815 cm tall. Rhizome shortly creeping to ascending, apex scaly; scales dark brown, narrowly triangular to subulate, entire to denticulate, at base fimbriate. Fronds caespitose; stipe shiny castaneous only at base, abruptly green toward rachis, 610 cm, 23 Χ as long as lamina, with unicellular glands or subglabrous, apex 2- or 3-forked; segments linear, 23 Χ 0.10.15(0.25) cm, base gradually decurrent onto stipe, minutely forked again at apex. Fronds without distinct rachis, green, other veins slender and subparallel to central axis, one veinlet per ultimate segment. Fronds herbaceous-leathery, grass-green. Sori 15 per segment, linear, 12 cm, on acroscopic veins and close to costa, at maturity covering entire surface; indusia brown, linear, membranous, free margin often with unicellular glands, entire-sinuate, opening toward main vein (costa), persistent but reflexed and covered by sporangia at maturity. Spores with lophate perispore, average exospore length 3944 ΅m. Plants sexual tetraploid: 2n = 144.
In crevices of non-calcareous (often granitic) rocks in open or partially shaded situations; 11004100 m. Shaanxi, Shanxi, Taiwan, Xinjiang, Xizang [Afghanistan, India, Kazakhstan, Kyrgyzstan, W Mongolia, Nepal, Pakistan, Russia, Tajikistan; NW Africa, SW Asia, Europe, North America].
The type specimen is from Europe.
Asplenium septentrionale is the autotetraploid that originated by chromosome doubling in its diploid ancestor A. caucasicum. In the absence of a chromosome count, it can best be distinguished by its larger mean exospore length (more than 37 ΅m) and its broader lamina.
Asplenium septentrionale often hybridizes with other Asplenium species in Europe wherever they co-occur; one of these, A. Χheufleri Reichardt (A. septentrionale Χ A. lovisii), was recently found in Xinjiang.
黑边铁角蕨 hei bian tie jiao jue
Plants 48 cm tall. Rhizome erect, short, apex scaly; scales blackish brown, iridescent, narrowly triangular to subulate, entire. Frond simple, tufted; stipe purplish black, shiny, 512(20) mm, terete, with scales when young, subglabrous when old; lamina elliptic to narrowly triangular, 36(8.5) Χ 1.21.7(2) cm, base broadly cuneate to truncate, margin entire to undulate and with narrow black edge, apex obtuse. Rachis or midrib distinct abaxially, black and shiny; veins obscure, oblique, 2(or 3)-forked. Fronds leathery, adaxially olive-green when dry, brown abaxially. Sori linear, 36 mm, medial between midrib and margin; indusia grayish brown, linear-elliptic, thinly papery, entire and with narrow purple edge, opening toward midrib, persistent. Spores with cristate-alate perispore.
* In limestone crevices, forests; 11001400 m. Guangdong, Guangxi, Guizhou, Hunan, Jiangxi.
The type is from Guizhou (Pingfa).
Asplenium speluncae is a particular and distinct species. Hybrids with A. normale were probably described from Hunan as A. Χxianqianense C. M. Zhang (Keys Vasc. Pl. Wuling Mount. 568. 1995, published as a species) (pro sp.) but are also known from Guangxi.
狹叶铁角蕨 xia ye tie jiao jue
Asplenium annamense Christ; A. pinfaense Christ.
Plants 2045(60) cm tall. Rhizome shortly creeping to erect, apex densely scaly; scales medium to dark brown, ovate-triangular, subentire to denticulate. Frond simple, ± clustered, subsessile or with stramineous stipe 15()7 cm; lamina linear to narrowly lanceolate, 1540(50) Χ 1.12(3) cm, usually more than 13 Χ longer than wide, gradually attenuate at both ends, base narrowly cuneate and decurrent on stipe, margin entire to repand or sinuate, with minute notches, apex long acuminate. Midrib (rachis) distinct, often slightly raised (semiterete) abaxially or adaxially; veins distinct, usually forked in their upper part. Frond leathery, brownish green to olivaceous when dry, with sparse minute brown triangular-stellate scales. Sori linear, 47 mm, starting close to midrib at an angle of (40°)50°70°(80°); indusium gray to yellowish brown, linear, relatively thick, entire, opening toward midrib, persistent. Spores with alate perispore.
On tree trunks or shaded wet rocks in forests; 13001600 m. Guangdong, Guangxi, Guizhou, Hainan, Yunnan [India, Malaysia, Myanmar, Thailand, Vietnam].
The type is from the Malay Peninsula (Perak).
Typical Asplenium scortechinii plants have narrow fronds, but intermediates with A. griffithianum exist. Plants from the Malay peninsula were found to be tetraploid (n = ca. 72) by Manton (in Holttum, Fl. Malaya 2: 625. 1954), but recent FCM results [??citation] show that, at least in China, this taxon is an aggregate of hexaploid (Hainan) and dodecaploid (Yunnan) species. In particular, the relationships with dodecaploid A. griffithianum needs further investigation.
Asplenium holophyllum Baker; A. iridiphyllum Hayata; A. nakanoanum Makino; A. scolopendrifrons Hayata.
Plants 1525(30) cm tall. Rhizome erect, short, apex densely scaly; scales blackish brown, narrowly triangular or ovate, margins denticulate. Fronds caespitose, simple; stipe stramineous, very short or absent, 0.42.5 cm; lamina lanceolate, 1525 Χ (1.5)1.92.5(3.9) cm, usually ca. 10 Χ longer than wide, base gradually decurrent and forming narrow wing on stipe, margin entire or sinuate in basal part, upward irregularly sinuate-crenate, and often notched, apex acuminate or acute. Midrib stramineous, not distinctly raised abaxially, adaxially raised; veins obscure, 1-forked. Fronds fleshy, grayish green when dry, with minute brown triangular to stellate scales, subglabrous when old. Sori brown, linear, 59(11) mm, starting close to midrib at an angle of (30°)45°50°(65°), and up to 2/3 width of lamina; indusia gray to yellowish brown, linear, entire, opening toward midrib, persistent. Spores with many perforations in outer subechinate-lophate perispore.
On tree trunks or wet rocks in forests; 1001600 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Sichuan, Taiwan, Xizang, Yunnan [Bhutan, India, Japan, Myanmar, Nepal, Vietnam].
The type is from NE India (Khasia).
Plants from Sikkim, India, were reported to be diploid hybrids by Bir ([??Current Science 29: 445447]. 1960) and by Mehra ([citation?]. 1961), but recent FCM results [??citation] show that at least the Guizhou plants are polyploids (1214x).
In its present circumscription this species includes different taxa that need further study. Asplenium Χxinyiense Ching & S. H. Wu (Bull. Bot. Res., Harbin 9(2): 82. 1989), a plant with abortive spores described from SW Guangdong and published as a species, is a hybrid between either A. griffithianum or A. loxogrammoides and A. wrightii (pro sp.). These species grow intermixed on cliffs near the type locality of A.Χxinyiense.
江南铁角蕨 jiang nan tie jue
Asplenium loxogrammoides Christ.
Plants 2040 cm tall. Rhizome erect, short, apex scaly; scales dark brown, narrowly triangular, entire. Fronds simple, clustered; stipe grayish stramineous, 24 cm, with scales; lamina lanceolate, 1530(36) Χ 2.53.5(4) cm, base decurrent on stipe, margin pale, ± entire to repand or sinuate, apex acuminate to subcaudate. Midrib (rachis) distinctly raised and semiterete on both surfaces in basal half of lamina, with dark brown to black filiform paleasters, brownish; veins obscure, 1-forked. Fronds herbaceous, dark green to brown when dry, subglabrous or with small stellate scales. Sori linear, 1.42.1(2.5) cm, often starting close to midrib or more medial, at an angle of (20°)30°40°, on acroscopic veinlets; indusia yellowish brown to deep brown, linear, thickly membranous, margin composed of thinly walled hyaline cells, entire to repand, opening toward midrib, persistent. Spores with alate perispore.
On rocks by streamsides in forests; 5001000 m. Guangdong, Guangxi, Guizhou, Hainan, Hubei, Hunan, Jiangxi, Sichuan, Taiwan, Yunnan [Vietnam].
The type is from Yunnan (Mengtze).
Asplenium holosorum is similar to A. ensiforme but generally grows at lower elevations. Its fronds are more herbaceous and their midrib (rachis) is flat near the apex but raised adaxially in the lower half of the frond and not sulcate. Both species have a similar epidermis structure with cryptopolocytic to basipolocytic (also called anomocytic, staurocytic, or cyclocytic) stomata. In both taxa, the outer free margin of the indusium consists of translucent cells with very thin walls, but this marginal rim is narrow (ca. 2 cell layers) in A. holosorum, but more elaborate and dentate-fimbriate in A. ensiforme.
No chromosome counts seem to be available but FCM results [??citation] suggest this taxon is probably hexaploid.
劍叶铁角蕨 jian ye tie jiao jue
Asplenium biscupe Hayata; A. ensiforme f. bicuspe (Hayata) Ching ex S. H. Wu; A. ensiforme var. bicuspe (Hayata) Tagawa; A. ensiforme var. parvum Tardieu & Ching; A. ensiforme f. stenophyllum (Beddome) Ching ex S. H. Wu; A. ensiforme var. stenophyllum (Beddome) Ching; A. gracilipes Ching & Y. X. Lin; A. melanolepis Baker (1890), not Franchet & Savatier (1879), nor Colenso (1888); A. tonkinense C. Christensen; A. stenophyllum Beddome.
Plants 3045(65) cm tall. Rhizome erect, short, apex scaly; scales dark brown to black, narrowly triangular, ca. 6 Χ 1 mm, entire. Fronds simple and clustered; stipe stramineous, 58(15) cm, base with scales, becoming subglabrous above; lamina narrowly lanceolate, 2040(50) Χ 1.52.5(4) cm, base decurrent on stipe, margin entire to repand, apex acuminate to caudate. Midrib stramineous, abaxially semiterete and raised, adaxially sulcate; veins obscure, 1-forked. Fronds leathery, yellowish green or brownish when dry, subglabrous. Sori linear, 1.32(3) cm, starting close to midrib at an angle of 15°25°(30°), on acroscopic veinlets; indusia yellowish brown or brownish green and becoming dark brown when dry, linear, papery, margin composed of thinly walled hyaline cells, fimbriate to entire, opening toward midrib, persistent. Spores without perforations in outer cristate-alate perispore. Plant sexual tetraploid: 2n = 144.
On rocks and tree trunks in forests; 8002800 m. Guangdong, Guangxi, Guizhou, Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan, Xizang, Yunnan [Bhutan, India, Japan, Myanmar, Nepal, Sri Lanka, Thailand, Vietnam].
The type is from Nepal.
Plants of Asplenium ensiforme with extremely narrow fronds, occurring at higher elevations, have sometimes been recognized as a separate species, variety, or form. A specimen from Taiwan with forked frond apices was described as A. biscupe, and we have seen such aberrant forked forms also from Yunnan. Forked fronds occur in all species of pteridophytes, often randomly in particular plants, and do not merit taxonomic status.
Plants from India were reported to be tetraploid (n = 72) by Mehra & Bir ([??Curr. Sci. 26: 151152]. 1957) and by Bir ([??Curr. Sci. 29: 445447]. 1960; [??Mem. Ind. Bot. Soc. 4: 116]. 1962; [??citation]. 1965). This species is similar to Asplenium holosorum; for differences, see below that species.
对开蕨 dui kai jue
Asplenium scolopendrium Linnaeus subsp. japonicum (Komarov) Rasbach, Reichstein & Viane; Phyllitis japonica Komarov.
Plants up to ca. 60 cm tall. Rhizome erect or ascending; scales brown, thin, narrowly triangular to triangular, entire. Fronds caespitose; stipe brown, 1020 cm, with sparse scales; lamina narrowly oblong, 1545 Χ 3.54.5(6) cm, leathery, base cordate, margin entire or slightly sinuate, apex acute. Midrib (rachis) distinct, brown basally and becoming green toward apex, raised and rounded on both sides, with small scales at base; lateral veins obscure, anadromous but running straight and almost parallel. Frond green and fleshy when fresh, after drying leathery and brownish green, abaxially subglabrous. Sori linear, usually 725 mm, on neighboring veinlets and opposite; indusia brown, linear, entire, opening toward each other (scolopendrioid), persistent. Spores with lophate (alate) perispore, average exospore length 3135 ΅m. Plants sexual tetraploid: 2n = 144.
Terrestrial; 7001000 m. Jilin (Changbai, Fusong, Jian), Taiwan [Japan, Korea, SE Russia; North America].
Due to its peculiar frond shape and double sori facing each other, Asplenium komarovii was often placed in the former satellite genus Phyllitis Hill. Modern studies confirm that it deserves no recognition. The group consists of two species: a diploid ancestral taxon (2n = 72) essentially in Europe, and a tetraploid in North America (including Mexico) and NE Asia (Mitui, [citation?]. 1966; [citation?]. 1968).
水鳖蕨 shui bei jue
Scolopendrium delavayi Franchet, Bull. Soc. Bot. France 32: 29. 1885; Phyllitis delavayi (Franchet) C. Christensen; Schaffneria delavayi (Franchet) Tardieu; Sinephropteris delavayi (Franchet) Mickel.
Plants up to 15 cm tall. Rhizome erect, short; scales black, narrowly triangular, margins sparsely toothed. Fronds simple, clustered; stipe chestnut-black to black, shiny, 310 cm, subglabrous, cylindrical but with an adaxial groove; lamina orbicular-reniform, usually 2.56 cm in diam., base cordate, margins entire to slightly sinuate, thin. Dark midrib distinct at base and running well into lamina, other veins usually obscure, veinlets mostly free but casually[some?] connected near margin. Fronds herbaceous or papery, brownish green or brown after drying, minutely hairy. Sori linear, in pairs facing each other, confluent at maturity; indusia brownish, linear, persistent. Spores with lophate (alate) perispore.
On shaded and wet rocks in forests; 6001800 m. S Gansu, S Guizhou, Sichuan, Yunnan [N India, N Myanmar, Nepal].
The type is from Yunnan.
Based on its peculiar morphology and scolopendrioid sori, Asplenium delavayi has been placed in various satellite genera; these have now been abandoned because modern studies show them all nested within Asplenium. See Mickel ([??Amer. Fern J. 66: 8182]. 1976) for a discussion on the differences with supposedly related groups.
扁柄巢蕨 bing chao jue
Asplenium longistipes (Ching ex S. H. Wu) Viane; Neottopteris humbertii (Tardieu) Tagawa; N. longistipes Ching ex S. H. Wu.
Plants up to 30 cm tall. Rhizome erect, short, scaly; scales brown, triangular-ovate, margins sparsely ciliate. Fronds caespitose; stipe greenish stramineous, 48 cm, soft, without wide wing on lateral side, base with dark brown scales or subglabrous; lamina spatulate, 1822 Χ 3.55 cm, lower part decurrent on stipe and cuneate, margin entire, apex cuspidate to shortly caudate. Midrib (rachis) almost flat to slightly semiterete, stramineous, abaxially occasionally with small brown scales; veins obscure adaxially, faintly visible abaxially, simple or forked. Fronds leathery, after drying grayish green, subglabrous. Sori linear, 11.5(2.5) cm, on acroscopic veinlets, occupying 1/22/3 of subtending vein; indusia grayish yellow, linear, thickly membranous, entire, persistent. Spores with lophate areolo-fenestrate perispore. Plants sexual hexaploid: 2n = 216.
Epilithic on limestone rocks in wet forests; 800900 m. Guangxi, Hainan, SE Yunnan [Laos, Thailand, Vietnam].
The type is from Vietnam.
Specimens of Asplenium humbertii from Hainan have longer sori than the type. The species is rather similar to A. grevillei Wallich ex Hooker & Greville from India and Thailand which differs by its smaller scales, distinctly keeled midrib, and areolo-fenestrate perispore (Wei & Dong, [??Nordic J. Bot. 30: 90103]. 2012). However, the relationship between A. grevillei, A. humbertii, and A. antrophyoides requires further study.
狭翅巢蕨 xia chi chao jue
Asplenium latipes (Ching ex S. H. Wu) Viane; A. subspathulatum Rosenstock; Neottopteris antrophyoides (Christ) Ching; N. latipes Ching ex S. H. Wu.
Plants 3060 cm tall. Rhizome erect and short, apex scaly; scales dark brown, narrowly triangular, entire. Fronds caespitose, subsessile or with extremely short stipe; stipe stramineous, soft; lamina spatulate to very narrowly obovate, 2560 Χ 4.56.5(8) cm, gradually reduced into a 12 cm wide lateral wing; margin entire, often cartilaginous, apex cuspidate to caudate. Midrib flat to carinate at base abaxially, stramineous, subglabrous but with dark brown lanceolate scales at its base; veins visible on both sides, simple or forked, parallel. Fronds subleathery, brownish green to dark green after drying, subglabrous. Sori restricted to upper part of lamina, linear, 1.52.5 cm, borne on acroscopic veinlets and occupying 2/33/4 of their length; indusia brownish, linear, membranous, entire, persistent. Spores with lophate (cristate) perispore. Plants sexual tetraploid: 2n = 144.
On limestone rocks or on tree trunks in dense forests; 3001300 m. Guangdong, Guangxi, Guizhou, Hunan, Sichuan, Yunnan [Laos, Thailand, Vietnam].
The type is from Guizhou.
An aberrant monstrosity, described as Asplenium antrophyoides var. grossidentatum Bonaparte ex Tardieu (Asplιn. Tonkin, 25. 1932; Neottopteris antrophyoides var. cristata Ching & S. H. Wu), with a lacerate frond, was collected in Vietnam and SE Yunnan[add to synonymy or is it nom. inval.??]. Another form with a very wide frond base and described as Neottopteris latipes by Ching is also treated as a synonym; the perispores are similar (Wei & Dong, [??Nordic J. Bot. 30: 90103]. 2012).
巢蕨 chao jue
Asplenium neohainanense Viane; Neottopteris hainanensis Ching; N. nidus (Linnaeus) J. Smith; N. rigida Fιe; N. salwinense Ching; Thamnopteris nidus (Linnaeus) C. Presl.
Plants 11.2 m tall. Rhizome erect, thick and short, woody, apex scaly; scales dark to purplish brown, narrowly triangular to linear-subulate, margin ciliate to fimbriate. Fronds caespitose; stipe pale brown, up to 5 cm, woody, when dry semiterete abaxially, base densely scaly; lamina lanceolate, 90120 Χ (8)915 cm, gradually decurrent on stipe, base cuneate, margin entire, apex acute to acuminate. Midrib raised and semiterete on upper adaxial side but flat abaxially, subglabrous, grayish to pale brown; veinlets simple or forked, parallel and connected at their apex to marginal vein. Fronds papery or thinly leathery, when dry grayish green, glabrous. Sori linear, 35 cm, on acroscopic side of veinlets, running from near their base up to 1/2 of their length; basal part of lamina usually sterile; indusia brownish, linear, thickly membranous, entire, persistent. Spores with lophate (costate to cristate) perispore. Plants sexual tetraploid: 2n = 144.
Clustered on tree trunks or rocks in rainforests; 1001900 m. ?Guangdong, Guangxi, ?Guizhou, Hainan, Taiwan, Xizang, Yunnan [Cambodia, India, Indonesia, Japan, Laos, Malaysia, Myanmar, Sri Lanka, Vietnam; tropical regions of E Africa and Australia, Pacific islands (Polynesia)].
The type is from Indonesia (Java).
Asplenium nidus is accepted here in a broad sense and constitutes a species complex (e.g., Murakami et al., [??in M. Kato, Biol. of Biodiversity, 5366]. 1999; Yatabe et al., [??Amer. J. Bot. 88: 15171522]. 2001). The variability of its frond and perispore morphology (e.g., Wei & Dong, [??Nordic J. Bot. 30: 90103]. 2012), as well as other phenetic characters, is not well studied in relation to its molecular diversity.
Asplenium setoi N. Murakami & Serizawa, recently described from Japan, might be present at low elevations in China; typical specimens can be distinguished from A. nidus by their keeled to boat-shaped midrib. Another species regularly confused with both A. setoi and A. nidus is the often cultivated A. australasicum (J. Smith) Hooker, a South Pacific taxon.
Based on their particular venation pattern, taxa resembling Asplenium nidus have been recognized as a separate section (A. sect. Thamnopteris Hooker & Baker), a subgenus (A. subgen. Thamnopteris Tardieu), or as a genus of its own (Neottopteris J. Smith; syn. Thamnopteris C. Presl). Plants are often epiphytes with their large simple fronds growing in a close spiral and forming the typical birds nest. Veins departing from the midrib (rachis) fork anadromously, run almost parallel and straight to the margin where they connect to a common submarginal vein. However, recent molecular studies do not support the separation of this group as a separate genus. The clade consists of ca. 1530 species, and modern research shows that more taxa may await description. A critical revision of the group is urgently needed. Members occur mainly in rainforests of tropical Asia and the Pacific. A few taxa are widely cultivated as house plants and sold as birds-nest fern. Many plants in commerce belong to A. australasicum, which can be distinguished from true A. nidus by its abaxially dark brown carinate midrib.
黑鳞巢蕨 hei lin chao jue or 阔足巢蕨 kuo zu chao jue
Asplenium latibasis (Ching) Viane (1991), not (Ching & Shing) Nakaike (1986); A. phyllitidis D. Don subsp. malesicum Holttum; A. subantiquum (Ching ex S. H. Wu) Viane; Neottopteris latibasis Ching; N. subantiqua Ching ex S. H. Wu.
Plants 60100 cm tall. Rhizome erect, dark brown, short and thick, woody, apex scaly; scales brown to dark brown, narrowly triangular, membranous, apex acuminate, fimbriate. Fronds caespitose; stipe dark brown, very short, 12 cm, when dry semiterete on both sides, base with scales similar to those on rhizome; lamina lanceolate to narrowly lanceolate, 5060(100) Χ (4)89(12) cm, gradually decurrent toward stipe, margin entire, with a cartilaginous narrow edge or slightly recurved when dry, apex acute. Midrib prominent on both sides, semiterete to obtusely carinate abaxially, grooved adaxially when dry, dark brown or grayish brown, glabrous; veinlets visible, forked or simple, parallel, at their apex united to marginal vein. Fronds firmly papery to subleathery, green but brownish when dry, glabrous. Sori linear, 23 cm, on acroscopic veinlets, running from near their base up to (1/2)2/33/4 of their length, lower part of lamina usually sterile; indusia brown to dark brown, linear, thickly membranous, entire, persistent. Spores with echinate perispore. Plants tetraploid, 2n = 144.
On tree trunks in dense forest understories; 1001100 m. Hainan, SE Yunnan [from Malaysia north to Thailand, Vietnam, Myanmar, and E India (Assam)[please just list individual countries, e.g., E India (Assam), Malaysia, Myanmar, Thailand, Vietnam]].
Asplenium oblanceolatum is similar to other members of the A. nidus complex but differs from all by its echinate perispores (Holttum, [??Gard. Bull. Singapore 27: 143154]. 1974, Zhang et al., [??citation]. 1976, Wei & Dong, [??Nordic J. Bot. 30: 90103]. 2012). In addition, it differs from small A. nidus by its midrib structure, and from A. phyllitidis and A. antiquum by its narrower scales.
大鳞巢蕨 da ling chao jue
Neottopteris antiqua (Makino) Masamune; N. rigida Fιe var. erubescens Nakai; Thamnopteris antiqua (Makino) Makino.
Plants 80100 cm tall. Rhizome erect, massive; scales on apex brown to dark grayish brown, ovate-triangular, entire to fibrillose. Fronds caespitose; stipe pale brown to dark brown, 25(7) cm, woody, terete abaxially, adaxially grooved, scales at stipe base numerous, similar to those on rhizome, 1530 Χ 26 mm; lamina narrowly lanceolate, 75100 Χ 6.58.5(15) cm, cuneate at base, margin entire and cartilaginous. Midrib raised on both sides, semiterete abaxially, dark brown, subglabrous; veins slightly raised on both sides, forked or simple, parallel, connected to marginal vein. Fronds leathery, after drying brownish green or brownish. Sori linear, 34 cm, on acroscopic veins, occupying 2/33/4 of subtending vein, lower parts of lamina usually sterile; indusia brownish or grayish brown, linear, thickly membranous, entire, persistent. Spores with lophate perispore. Plants tetraploid, 2n = 144.
On rocks or tree trunks in forests; ca. 900 m. Fujian, ?Hunan, Taiwan [Japan, Korea].
Asplenium antiquum was wrongly reported from Hainan (Wu, FRPS 4(2): 137. 1999). The type is from S Japan.
长叶巢蕨 chang ye chao jue
Type: Nepal, Wallich s.n. (lectotype, designated here, BM).
Asplenium colaniae Tardieu; A. nidus Linnaeus var. phyllitidis (D. Don) Beddome; A. nidus var. simonsianum (Hooker) Christ; A. simonsianum Hooker; Neottopteris phyllitidis (D. Don) J. Smith; N. simonsiana (Hooker) J. Smith; Thamnopteris orientalis C. Presl; T. phyllitidis (D. Don) C. Presl; T. simonsiana (Hooker) T. Moore.
Plants (50)70110 cm tall. Rhizome erect, thick and short, woody, apex with scales; scales brown to blackish brown, triangular to ovate, membranous, entire or fimbriate. Fronds caespitose; stipe pale brown to grayish brown, 37 cm, woody, abaxially terete, broadly grooved adaxially when dry, glabrous, with two lateral ridges, base with scales similar to those on rhizome; lamina narrowly lanceolate to lanceolate, (48)70105 Χ 58 cm, widest at middle and gradually decurrent toward base, margin entire, apex acute to caudate. Midrib semiterete and grayish to yellowish pale brown abaxially, broadly grooved and pale, yellowish brown adaxially when dry, glabrous; veins faintly visible, forked, or simple. Fronds papery or thinly leathery, brownish green when dry, glabrous. Sori linear, 23 cm, on acroscopic veinlets, from close to midrib to 1/23/4 as long as subtending veinlet, lower part of lamina usually sterile; indusia brownish, linear, membranous, entire, persistent. Spores with lophate (areolo-fenestrate) perispore. Plants tetraploid, 2n = 144.
On rocks or tree trunks in forests by streamsides; 6001400 m. Guangxi, Guizhou, Hainan, Sichuan, Xizang, Yunnan [Bangladesh, Bhutan, India, Indonesia, Malaysia, Myanmar, Nepal, New Guinea, Thailand].
The type is from Nepal.
Asplenium simonsianum and A. colaniae are reduced to synonymy. Their types have a lamina 35 cm wide; we consider them extremely narrow forms of A. phyllitidis. The perispores of A. phyllitidis, A. simonsianum, and A. colaniae are closely similar and areolo-fenestrate.
过山蕨 guo shan jue
Antigramma sibirica J. Smith; Camptosorus sibiricus Ruprecht; Phyllitis sibirica (Ruprecht) Kuntze; Scolopendrium sibiricum (Ruprecht) Hooker.
Plants up to 20 cm tall. Rhizome erect; scales blackish brown, narrowly triangular, entire. Fronds caespitose; dimorphic, sterile fronds smaller with stipe 13 cm and lamina 12 Χ 0.50.8 cm, elliptic, base broadly cuneate and slightly decurrent, apex obtuse; fertile fronds larger with stipe 15 cm, lamina lanceolate, 1015 Χ 0.51 cm, base cuneate (cuneate-attenuate), decurrent, margin entire or slightly sinuate, apex acuminate, flagelliform (38 cm) and terminating in a gemma or rooting. Veins anastomosing, obscure, with 13 rows of areolae, usually one row of areolae close to midrib (rachis), other veinlets free. Fronds herbaceous, dark green after drying, subglabrous. Sori in 13 irregular rows on both sides of midrib (rachis), linear to elliptic; indusia gray-green to brownish, linear or elliptic. Spores with lophate (cristate-alate) perispore. Plants sexual diploid: 2n = 72.
On rocks in forests; 3002000 m. Hebei, Heilongjiang, Henan, Hubei, Jiangsu, Jilin, Liaoning, Nei Mongol, Ningxia, Shaanxi, Shandong, Shanxi, Sichuan [Japan, Korea, Russia].
The type is from Siberia.
Based on its peculiar morphology and venation pattern, Asplenium ruprechtii has often been put in the satellite genus Camptosorus Link, containing two species (one in Asia, the other in North America). Modern studies have shown that Camptosorus nests within Asplenium and does not form a separate clade.
Asplenium ruprechtii is diploid (Kurita, [citation?]. 1965; Mitui, [citation?]. 1965) and hybridizes with tetraploid A. anogrammoides (sarellii sensu Kurita et auct. Jap.) to form the sterile triploid A. Χkitazawae Sa. Kurata. It is also ancestral to the following species.
东海铁角蕨 dong hai tie jiao jue
Asplenium kobayashii Tagawa; ΧAsplenosorus castaneoviridis (Baker) Nakaike.
Plants 820 cm tall. Rhizome erect, short, scaly; scales black, narrowly triangular. Fronds caespitose; stipe adaxially green, abaxially shiny castaneous, subglabrous; lamina pinnatisect, dimorphic and variable; stipe of small fronds 24 cm, of large fronds 68 cm, lamina of small fronds narrowly triangular to linear, 59 Χ ca. 1 cm; lamina of large fronds ovate-lanceolate, 1114 Χ 25 cm, attenuate to both ends, apex acuminate; pinnae 79 (small fronds) or 1015 pairs, opposite or subopposite, sessile, lower segments of large fronds reduced and gradually more triangular-deltoid, middle segments narrowly triangular, 12 Χ 0.30.5 cm, base asymmetrical and adnate to rachis, acroscopic side (sub)auriculate, basiscopic side decurrent into narrow wing along rachis, margin repand to sinuate, apex obtuse; segments of small fronds elliptic to obovate, lowest segments largest, 58 Χ 0.30.5 cm, adnate to rachis, margin serrate, apex obtuse. Veins anadromously pinnate, obscure, veins simple or forked ending in submarginal hydathode. Fronds herbaceous, green to yellowish green when dry; rachis green, (sub)glabrous. Sori 36 per pinna, linear-elliptic, ca. 2 mm, on acroscopic veinlets; indusia green, linear-elliptic, entire, opening toward costa.
On rocky cliffs. Jiangsu, Liaoning, Shandong [Japan, Korea].
The type is from China (Shandong, Zhi-fu).
We consider Asplenium castaneo-viride to be a good species, not a sterile hybrid (Ching & Iwatsuki, [??J. Jap. Bot. 57:129132]. 1982), because Lovis et al. ([??Brit. Fern Gaz. 10: ??]. 1972) and Lin and Sleep ([??in K. H. Shing & K. U. Kramer (eds.), Proc. Int. Symp. Syst. Pterid. 111127]. 1989) have shown that this is a fertile allotetraploid, which originated via chromosome doubling in the sterile hybrid between A. ruprechtii and A. incisum.
苍山蕨 cang shan jue
Asplenium alternans Wallich, nom. nud.[Wallich ex T. Moore, nom. illeg. superfl.? or Wallich ex Hooker, nom. illeg. superfl.? Moore (Ind. Fil. 112. 1857) didnt provide a description for Wallichs name but cited A. dalhousiae in syn. Hooker (Sp. Fil. 3: 92. 1860) provided description and also cited A. dalhousiae in syn.]; Ceterach alternans Kuhn & Decken; C. dalhousiae (Hooker) C. Christensen; Ceterachopsis dalhousiae (Hooker) Ching.
Plants 1315(25) cm tall. Rhizome erect, short; scales brown to dark brown, narrowly triangular to lanceolate, margin with short teeth. Fronds caespitose; stipe short, less than 2 cm; lamina narrowly elliptical-obovate, 415 Χ 1.85 cm, gradually narrowed at base, pinnatipartite, apex acute to obtuse; segments 1014 pairs, alternate, ovate to narrowly ovate, 812 Χ 58 mm, margin entire and hyaline, apex obtuse. Veins obscure, nothocatadromous (anadromous base pattern but several middle pinnae with their basal vein pair catadromous), lateral veins 68 pairs, often forking near margin. Fronds firmly papery, brownish yellow to dark brownish green when dry, glabrous, average guard cell length 4554 ΅m. Sori 57 pairs per segment, 35 mm, oblique, on acroscopic secondary veinlets; indusia pale brown, linear, membranous, mostly opening toward costa. Spores pale brown, average exospore length 2630 ΅m, outer perispore folded (lophate) and with small pores. Plants sexual diploid: 2n = 72.
On rocks; 10003000 m. ?Xizang [NW Himalaya from Afghanistan to Nepal[please just list individual countries]; isolated populations in Africa, North America (Mexico, United States)].
Asplenium dalhousiae is a relatively common W Himalayan fern expected to occur in Xizang and is recorded provisionally in the Flora. This and the following three taxa (A. paucivenosum, A. magnificum, A. qiujiangense) were often included in the genus Ceterachopsis (J. Smith) Ching (Bull. Fan Mem. Inst. Biol. Bot. 10(1): 8. 1940; 苍山蕨属 cang shan jue) on the basis of their peculiar frond morphology. The group consists of 4 or 5 species, mainly from the subtropical mountains of SW China, and one (A. dalhousiae) more widely distributed taxon. Recent studies have confirmed that this clade does not merit generic rank.
疏羽苍山蕨 shu mai cang jue
Asplenium paucivenosum f. minus Bir; Ceterach paucivenosa Ching; Ceterachopsis paucivenosa (Ching) Ching.
Plants 1525 cm tall. Rhizome erect, short; scales brown, triangular, margins entire or sparsely dentate. Fronds herbaceous, clustered; stipe short; lamina narrowly elliptical-obovate, 1116 Χ 1.56 cm, gradually narrowed to base, pinnatipartite, apex acute; segments 613 pairs, alternate, perpendicular to midrib (rachis), basal segments semicircular, upper ones oblong to triangular-ovate, apex obtuse, middle segments 12.5 Χ 11.5 cm, entire or slightly sinuate, glaucous or green becoming brown with distinct hyaline narrow margin after drying. Veins faintly visible, nothocatadromous (anadromous base pattern but several middle pinnae with their basal vein pair catadromous), average guard cell length 7993 ΅m. Sori short, subelliptic to ensiform-semilunulate, 28 Χ 12 mm, oblique, usually 2 or 3 pairs per segment, in middle between costa and margin, on acroscopic secondary veinlets; indusia grayish brown, semi-elliptic to ensiform-semilunulate, thinly membranous, entire, persistent. Spores with lophate-reticulate perispore, average exospore length 4551 ΅m. Plants sexual tetraploid: 2n = 144.
On rocky cliffs in mixed forests by streamsides; 20002700 m. Xizang, Yunnan [Himalaya from India to Bhutan[please just list individual countries]].
The type is from Yunnan.
Asplenium paucivenosum is a tetraploid species similar to the diploid A. dalhousiae but has much larger spores (and stomates) with a mean exospore length over 40 ΅m (less than 35 ΅m in A. dalhousiae) and a different, more fenestrate outer perispore. According to Nakaike (Ferns of Nepal 2: 45, 1986), it was confused with A. dalhousiae by Itτ (in H. Hara, Fl. E. Himal. 1: 487. 1966; Fl. E. Himal. 2: 214. 1971) and Iwatsuki (in Ohashi, Fl. E. Himal. 3: 194. 1975). It is even more easily confused with A. magnificum which has a larger mean exospore length (48 ΅m in A. paucivenosum versus 52 ΅m in A. magnificum). In critical cases, a chromosome count is needed for identification.
大叶苍山蕨 da ye cang shan jue
Ceterachopsis magnifica Ching, Bull. Fan Mem. Inst. Biol. Bot. 11: 56. 1941; Asplenium paucivenosum (Ching) Bir f. majus Bir.
Plants 1535 cm tall. Rhizome erect, short; scales brown to reddish brown, triangular. Fronds clustered; stipe brown, 24 cm, with brown triangular scales similar to those on rhizome, becoming subglabrous when old; lamina narrowly elliptical to elliptical, (13)1932 Χ 45 cm, attenuate to both ends, base cuneate, pinnatipartite, apex subobtuse; segments 1318 pairs, alternate, subperpendicular to midrib (rachis), basal segments semicircular, upper ones oblong to narrowly triangular-ovate, apex obtuse, middle ones (1.2)22.5 Χ (1)1.31.6 cm, margin hyaline and entire to slightly sinuate. Veins obscure, nothocatadromous (anadromous base pattern but several middle pinnae with their basal vein pair catadromous). Fronds subleathery, brownish to grayish green, with minute scales, average guard cell length 8499 ΅m. Sori 4 or 5 pairs per segment, elliptic, short, 35(8) mm, median on acroscopic secondary veinlets in middle between costa and margin; indusia brown, semi-elliptic, membranous, entire, persistent. Spores with lophate-reticulate perispore, average exospore length 5058 ΅m. Plants octoploid: 2n = 288*.
* On rocks under mixed forests; ca. 2800 m. W Yunnan (Yangbi).
Asplenium magnificum is easily confused with A. paucivenosum (see above), but in general, it is a larger plant. In critical cases, a chromosome count will be necessarily.
俅江苍山蕨 qiu jiang cang shan jue
Asplenium dulongjiangense Viane, nom. superfl.; A. latibasis (Ching & Shing) Nakaike; A. latilobum Viane, nom. superfl.; Ceterachopsis latibasis Ching & Shing; C. qiujiangensis Ching & S. H. Fu.
Plants up to 40 cm tall. Rhizome erect, short; apex scales brown, triangular, entire. Fronds caespitose; stipe short, 0.51.5(2) cm, winged; lamina narrowly elliptical (lanceolate), (15)2030(35) Χ 46(8) cm, gradually narrowed to base, pinnatipartite, apex acute; segments (11)1417 pairs, alternate, perpendicular-oblique to midrib (rachis) and usually falcate, triangular to narrowly triangular, middle segments 35 Χ 11.5(2) cm, margin hyaline and entire, usually with a gemma near pinna base at acroscopic margin, apex obtuse to acute. Veins obscure, nothocatadromous (anadromous base pattern but several middle pinnae with their basal vein pair catadromous). Fronds firm, gray-brown after drying, subglabrous, rachis abaxially with sparse small scales, average guard cell length 7489 ΅m. Sori 35 pairs per segment, linear, up to 1 cm, median on acroscopic secondary veinlets in middle between costa and margin; indusia brown, linear, at maturity obscured by sporangia. Spores with reticulate perispore with large pores and faint crests, average exospore length 4145 ΅m.
* On rocks in forests; 18002500 m. NW Yunnan.
Asplenium qiujiangense is similar to A. magnificum but has more falcate and more acute segments, with a gemma at the acroscopic margin at the pinna base. It can also be confused with A. paucivenosum, which has a larger mean exospore length (48 ΅m in A. paucivenosum versus 43 ΅m in A. qiujiangense). In critical cases, a chromosome count will be necessarily.
Ceterachopsis chiukiangensis Ching & S. H. Fu and C. latiloba Ching & K. H. Shing (in Y. L. Chang et al., Sporae Pterid. Sin. 256. 1976) are conspecific with Asplenium quijiangense, but the two were not validly published because no Latin description or diagnosis was provided (Vienna Code, Art. 36.1).
药蕨 yao jue
Ceterach officinarum Willdenow; Hemidictyum ceterach Beddome.
Plants up to 12 cm tall. Rhizome erect, short; scales dark brown in center to pale brown at margin, ovate-triangular, margin sparsely toothed. Fronds caespitose; stipe brown, short, 24 cm; lamina narrowly elliptic, 58 Χ 11.6 cm, reduced to base, pinnatipartite, apex obtuse; segments 68 pairs, alternate, triangular, bases adnate to rachis, entire or slightly sinuate, apex obtuse. Veins obscure, nothocatadromous (anadromous base pattern but several middle pinnae with their basal vein pair catadromous), anastomosing and forming subhexagonous submarginal areolae. Fronds subleathery, dark green to brownish green after drying, adaxially subglabrous, abaxially densely covered with brownish, ovate-lanceolate to ovate scales, average stomatal guard cell length 4048 ΅m. Sori linear, usually on acroscopic secondary veinlets and in middle between costa and margin; indusia rudimentary. Spores with lophate (costate) perispore, average exospore length 3742 ΅m. Plant sexual tetraploid: 2n = 144.
In dry rock crevices; 14002600 m. Xinjiang, N Xizang [W Himalaya to N Africa, Europe[please list individual countries adjacent to China first and then generalized range for distribution extending beyond these countries, e.g., Afghanistan, India, Kashmir; N Africa, SW Asia, Europe]].
Asplenium ceterach is widely distributed in S Europe and was often put into the separate genus Ceterach Willdenow (Anleit. Selbststud. Bot. 578. 1804, nom. cons.; 药蕨属 yao jue shu) on the basis of its peculiar pinnatipartite fronds, submarginally anastomosing veins, and the dense, abaxial scale covering. However, all these characters also occur in other clades within the family, and molecular studies (Van den Heede et al., [??Amer. J. Bot. 90: 481495]. 2003) have shown that Ceterach consists of two separate clades both nested within Aspleniaceae and not meriting generic status. This species is autotetraploid, its diploid ancestor (A. javorkeanum Vida), from which it can be distinguished by its larger spores, is only known from the Mediterranean area.
欧亚铁角蕨 ou ya tie jiao jue
Asplenium ramosum Linnaeus, nom. rej.; A. trichomanes-ramosum Linnaeus, nom. rej.
Plants small, 815 cm tall. Rhizome erect or ascending, short, apex scaly; scales black, narrowly triangular, often with glandular appendages at base. Fronds caespitose; stipe reddish brown or castaneous only at base, upward green, 26 cm, with dark brown narrow scales or subglabrous; lamina linear, 615 Χ 11.3 cm, base reduced, apex acute, 1-pinnate; pinnae 1420 pairs, rhombic-ovate to orbicular, stalked, basal pinnae opposite, upper pinnae alternate, middle pinnae 46 Χ 46 mm, margins crenate to (sometimes) deeply incised, apex obtuse, lower 2 or 3 pairs reduced and gradually deltoid, base subtruncate. Vein pattern obscure, anadromously pinnate, veins simple or forked, not reaching margin. Fronds herbaceous, green; rachis green, adaxially longitudinally grooved. Sori 48 per segment, linear, 11.5 mm, confluent at maturity; indusia greenish white, semi-elliptic, membranous, entire or slightly toothed, opening toward costa and persistent. Spores with average exospore length 3033 ΅m, perispore lophate (alate). Plants sexual diploid: 2n = 72.
In crevices of calcareous, dolomitic, or serpentine rocks; 39004500 m. Sichuan, Taiwan, Xinjiang, Xizang [Afghanistan, Himalaya[please list countries], India, Japan, Nepal, Pakistan, Russia; Europe, North America].
三翅铁角蕨 san chi tie jiao jue
Asplenium anceps Lowe ex Hooker & Greville[although IPNI lists Buch (Phys. Beschr. Canar. Ins. 189. 1825), only the name appears on that page; a full description and the ID credited to Lowe is found in Hook. & Grev.] var. proliferum Nakai; A. trialatum C. Christensen; A. trichomanes Linnaeus var. centrochinense Christ.
Plants 1530 cm tall. Rhizome erect, short, apex scaly; scales narrowly triangular, 3.5(5) Χ ca. 0.5 mm, with brown central stripe and paler clathrate borders, entire. Fronds caespitose; stipe shiny brown to black, 35 cm, stout, base terete, densely scaly, upward subglabrous and becoming trigonous and then triangular, abaxially carinate to winged and adaxially flat to sulcate with a brown membranous wing along each lateral side, stipe and rachis usually persistent after shedding of pinnae; lamina linear, 1228 Χ 12.5 cm, attenuate to both ends, 1-pinnate; pinnae 2035 pairs, (sub)opposite to alternate, usually at right angles to rachis, sessile, middle pinnae oblong to rectangular, 513 Χ 27 cm, base asymmetrical, acroscopic side truncate, sometimes auriculate, parallel with or covering rachis, basiscopic side narrowly cuneate, margin crenate, apex obtuse; basal pairs of pinnae gradually reduced, flabelliform. Veins pinnate, costa obscure, veins 1-forked. Fronds papery, green or brownish green when dry; rachis stout, shiny dark brown to black, subglabrous, triangular, abaxially winged, adaxially sulcate with two entire, lateral, brown membranous wings, often gemmiferous near apex, with 1 or 2(or 3) scaly axillary buds. Sori 36(11) per pinna, linear, 12 mm; indusium grayish green, semi-elliptic, membranous, entire, opening toward costa.
On wet rocks or in acidic soil in forests; 4001400 m. Anhui, Fujian, Gansu, Guizhou, Henan, Hubei, Hunan, Jiangxi, Shaanxi, Sichuan, Taiwan, Yunnan, Zhejiang [Japan, Korea, Myanmar].
The type is from Japan.
As presently circumscribed, Asplenium tripteropus is an aggregate of a diploid and a tetraploid species (2n = 72 or 144) both in China and in Japan (Mitui, [citation?]. 1965; Tatuno & Kawakami, [citation?]. 1969; Wang, [citation?]. 1989). Further studies will need to characterize and distinguish them, not only from each other but also from the non-gemmiferous, Macaronesian A. anceps.
This species is similar in outline to Asplenium trichomanes, but the plants can easily be distinguished by their triangular and winged stipe and rachis, and by the presence of a bud on some fronds.
铁角蕨 tie jiao jue
Asplenium densum Brackenridge, p.p.; A. melanolepis Colenso, p.p. (1888), not Franchet & Savatier (1879).
Plants 1030 cm tall. Rhizome erect, short, scaly; scales narrowly triangular, 3(4) Χ ca. 0.5 mm, with opaque, red to dark brown central stripe and paler narrow clathrate borders, entire. Fronds caespitose; stipe shiny castaneous-brown, 28 cm, base scaly, upward subglabrous, abaxially semiterete but adaxially grooved, with brown, membranous and subentire narrow wings, texture papery, stipe and rachis usually persisting after shedding of pinnae; lamina linear, 1025 Χ 0.91.6 cm, base slightly reduced, 1-pinnate, apex acute and 24 mm wide; pinnae ca. 2030 pairs, usually obliquely inserted, sessile, middle pinnae elliptic or ovate to orbicular, 2.57.5 Χ 24 mm, base subsymmetrical, cuneate, margin crenate, apex obtuse; lower pinnae gradually reduced. Veins pinnate, costa obscure, veins obliquely simple or up to 2-forked, basal acroscopic vein usually 2-forked. Fronds papery, green or brown when dry, average stomatal guard cell length 3542 ΅m; rachis castaneous, shiny, subglabrous, abaxially terete, adaxially grooved and with a relatively low lateral brown membranous wing. Sori 48 per pinna, oval to linear, 13.5 mm, usually on acroscopic vein; indusium white or brown after drying, oval to linear, membranous, free margin repand to entire, opening toward costa, persistent. Spores with lophate perispore, average exospore length 2731 ΅m. Plants sexual diploid: 2n = 72.
In crevices of non-calcareous (sandstone, granite) rocks, in open or half-shaded situations; 4003400 m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi, Shanxi, Sichuan, Taiwan, Xinjiang, Xizang, Yunnan, Zhejiang [worldwide in all temperate zones, in tropics on high mountains].
The type is from Europe.
Asplenium trichomanes is often circumscribed as a collective species consisting of various cytotypes (six taxa are known from C and S Europe). These taxa, often recognized for convenience at the subspecific level, are reproductively isolated (producing sterile hybrids when growing together) and are treated as species in this flora. The species can be distinguished by microcharacters, by flow cytometry or by counting chromosomes. When the complex is subdivided into species or subspecies, diploid silicolous cytotypes have been designated (since Lovis, [??Brit. Fern Gaz. 10: 1320]. 1964) as A. trichomanes s.s. or as subsp. trichomanes; the tetraploid as A. lovisii or as subsp. quadrivalens D. E. Meyer. A lectotype was selected in this sense by Viane (in Jonsell & Jarvis, [??Nordic J. Bot. 14: ??]. 1994), but as a result of the Linnean typification project this selection was overruled by Grubov ([citation?]. 1984), who selected the tetraploid specimen in herb LINN as type. However, pending a proposition to conserve the name A. trichomanes with a conserved type, based on a diploid plant, we use the name A. trichomanes s.s. in its traditional (2x) sense in compliance with ICBN Art. 57.1.
Asplenium Χlusaticum D. E. Meyer (the sterile hybrid between A. trichomanes and A. lovisii) is not rare where both parents grow together. It can be easily recognized by its aborted spores. In China, this hybrid is cytologically confirmed for Sichuan (Longchi).
铁角蕨 tie jiao jue
Asplenium densum Brackenridge, p.p.; A. melanolepis Colenso, p.p. (1888), not Franchet & Savatier (1879); A. quadrivalens (D. E. Meyer) Landolt; A. trichomanes Linnaeus subsp. quadrivalens D. E. Meyer.
Plants 1030 cm tall. Rhizome erect, short, scaly; scales narrowly triangular, 3.5(5) Χ ca. 0.5 mm, with opaque dark brown to black central stripe and paler narrow clathrate borders, entire. Fronds caespitose; stipe shiny castaneous or dark red-brown to black, 28 cm, base scaly, upward subglabrous, abaxially semiterete but adaxially sulcate, with brown, membranous and subentire narrow wings, texture papery, stipe and rachis usually not very long persistent after shedding of pinnae; lamina linear, 1025 Χ 0.91.6 cm, base slightly reduced, 1-pinnate, apex acute and 25 mm wide; pinnae ca. 2030 pairs, usually at right angles to rachis, sessile, middle pinnae oblong to rectangular, 46(12) Χ 24(5) mm, base asymmetrical, truncate, acroscopic side occasionally auriculate, margin crenate, apex obtuse; lower pinnae gradually reduced, flabellate or triangular. Veins pinnate, costa obscure, veins oblique, simple or up to 2-forked, basal acroscopic vein usually 2-forked. Fronds papery, green or brown when dry, stomatal guard cells 4148 ΅m; rachis castaneous, shiny, subglabrous, abaxially terete, adaxially grooved and with a relatively high lateral brown membranous wing. Sori 48 per pinna, oval to linear, 13.5 mm, usually on acroscopic vein; indusium white or brown after drying, oval to linear, membranous, free margin repand to entire, opening toward costa, persistent. Spores with lophate (costate) perispore, average exospore length 3237 ΅m. Plants sexual tetraploid: 2n = 144.
In crevices of various kinds of rocks (often calcareous), in open or half-shaded situations; 4003400 m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi, Shanxi, Sichuan, Taiwan, Xinjiang, Xizang, Yunnan, Zhejiang [worldwide in all temperate zones, in tropics on high mountains].
The type is from Europe.
Tetraploid Asplenium lovisii has a broader ecological amplitude, i.e., it is not limited to non-calcareous substrates, and has a wider range than A. trichomanes.
Asplenium Χheufleri Reichardt (the tetraploid sterile hybrid between A. lovisii and A. septentrionale) was recently found in Xinjiang. The authors have not seen any specimens of A. Χalternifolium Wulfen (the triploid sterile hybrid between A. trichomanes and A. septentrionale).
滇南铁角蕨 dian nan tie jiao jue
Plants 1520 cm tall. Rhizome erect, short, scaly; scales dark brown, narrowly triangular, ca. 2 mm. Fronds caespitose; stipe shiny or dull blackish purple, 23 cm, abaxially terete, adaxially flat with lateral low ridge of distinctly separate fingerlike (digitiform) projections; lamina linear-lanceolate, 1217 Χ 11.3 cm, attenuate to both ends, 1-pinnate; pinnae 2532 pairs, opposite or subopposite, sessile and often deciduous, middle pinnae elliptic-triangular to oblong, ca. 7 Χ 5 mm, base asymmetrical, basiscopic side narrowly cuneate, acroscopic side truncate and parallel with rachis, often auriculate, margin crenate-sinuate to entire, apex obtuse; lower pinnae gradually reduced, becoming rhomboid-flabellate. Costa obscure, venation anadromously pinnate, with few 1(or 2)-forked or simple veins. Fronds subleathery, grayish green when dry; rachis blackish purple to blackish brown, shiny, abaxially terete, upper part of adaxial side shallowly sulcate and with 2 rows of relatively closely set fingerlike projections forming low ridge (often eroded and not easily visible on older fronds), occasionally with scaly bud in axil of lowest pinna. Sori 27 per pinna, linear or linear-elliptic, 12.5 mm, medial on subtending vein; indusia grayish brown, linear-elliptic, membranous, entire-sinuate, opening toward costa. Spores with lophate perispore, average exospore length 3135 ΅m.
* On rocks in forests; ca. 2000 m. Yunnan.
Asplenium microtum is similar to A. trichomanes and A. lovisii but clearly differs by the fingerlike papillae bordering the wings on the rachis. A similar wing structure is also found in A. glanduliserratum and A. humistratum, from which it differs by its gemmiferous rachis. However, not all plants have gemmae, and these species are kept separate, pending future research on this complex. The Mexican Asplenium hallbergii Mickel & Beitel, which has a similar rachis but with papillae more perpendicular to the wing, is perhaps distantly related.
肾羽铁角蕨 shen yu tie jiao jue
Plants (4)1023 cm tall. Rhizome erect, short, apex scaly; scales narrowly triangular, with opaque dark brown central stripe and paler clathrate borders, entire. Fronds caespitose; stipe blackish purple to black, shiny, (0.3)23 cm, abaxially terete, adaxially flat but laterally with fingerlike projections; lamina linear, (4)1220 Χ 0.81.2 cm, apex obtuse with terminal deltoid segment, reduced at base, 1-pinnate; pinnae (20)2842 pairs, opposite, subopposite, or alternate, sessile, middle pinnae oblong, (3)56 Χ 1.54 mm, base asymmetrical, acroscopic side truncate and parallel with rachis, basiscopic side narrowly cuneate, margin entire or slightly sinuate, apex obtuse; basal pinnae slightly deflexed and gradually reduced, orbicular to ovate. Costa obscure, venation anadromously pinnate, with few 1(or 2)-forked or simple veins, basal acroscopic vein usually 2- or 3-forked. Fronds subleathery, grayish green to dark green when dry; epidermis with uniseriate glandular hairs and average guard cell length 4655 ΅m; rachis black, shiny, abaxially terete, adaxially flat but laterally with dense fingerlike projections gradually united at their base into a low ridge. Sori brown, linear-elliptic, (0.6)11.5 mm, median to distal on subtending vein, (1 or)25 in apical part of pinna; indusia grayish brown or brownish green, semi-elliptic, membranous, sinuate-entire, opening toward costa. Spores with lophate perispore, average exospore length 3034 ΅m.
* On limestone cliffs in forests; 8002200 m. Guizhou, Hubei, Hunan, Sichuan, Yunnan.
Asplenium humistratum is similar to A. trichomanes and A. lovisii but clearly differs by the fingerlike papillae bordering the wings on the rachis. It is more similar to A. glanduliserrulatum, from which it differs by its larger size and by the shape of its basal pinnae.
腺齿铁角蕨 xian chi tie jiao jue
Plants 1012 cm tall. Rhizome erect, short, densely scaly; scales narrowly triangular, with opaque dark brown-black central stripe and paler clathrate border, entire. Fronds caespitose; stipe purplish black, shiny, 511 mm, abaxially terete, adaxially flat but laterally with separate reddish brown fingerlike projections; lamina linear, 612(14) Χ 0.91.2 cm, apex obtuse with terminal deltoid segment, 1-pinnate; pinnae 2030 pairs, (sub)opposite to alternate, sessile, middle pinnae elliptic to trapeziform-oblong, 46 Χ 23 mm, base asymmetrical, acroscopic side truncate to broadly cuneate, basiscopic side narrowly cuneate, margin entire to repand, apex obtuse; apical pinnae hardly reduced, lower pinnae slightly and gradually reduced, often hastate or becoming subdeltoid. Costa obscure, venation anadromously pinnate, veins 2 or 3 pairs, simple. Fronds herbaceous to subleathery, grayish green when dry; rachis purplish black, shiny, abaxially terete, adaxially flat but laterally with fingerlike projections, these separated and rarely coalescent at their base into a low ridge. Sori 25 per pinna, linear-elliptic, ca. 1 mm, median on subtending vein, located in apical part of pinnae; indusia grayish brown, semi-elliptic, entire, opening toward costa.
* On rocks in forests; 12002400 m. Guizhou, NW Yunnan.
Asplenium glanduliserrulatum differs from A. trichomanes and A. lovisii by the fingerlike papillae bordering the wings on the rachis. It differs from A. humistratum by its smaller size and by the shape of its basal pinnae.
江苏铁角蕨 jiang su tie jiao jue
Asplenium gulingense Ching & S. H. Wu; A. hangzhouense Ching & C. F. Zhang; A. parviusculum Ching.
Plants 612 cm tall. Rhizome erect, short, apex scaly; scales narrowly triangular to linear-subulate, with broad opaque black central stripe and narrow paler clathrate borders, margin with uniseriate hairs at scale base. Fronds caespitose; stipe castaneous to dark brown, shiny, terete, 13.5 cm, with brownish, hyaline fibrillose small scales (paleasters), subglabrous when old; lamina linear, 310 Χ ca. 1 cm, apex acute, 1-pinnate; pinnae 820(22) pairs, lower ones subopposite, hardly reduced, middle pinnae spreading horizontally, elliptic to trapeziform-oblong, 45 Χ 45 mm, base asymmetrical, acroscopic side truncate and close to rachis, basiscopic side narrowly cuneate, shortly stalked to subsessile, margin entire to sinuate, apex obtuse. Venation anadromously pinnate or with first basiscopic vein lacking, costa with 2 or 3 acroscopic veins, obscure, veins simple or 1-forked. Fronds papery, grayish green when dry; rachis castaneous to dark brown, shiny, with small scales, semiterete and adaxially flat or with two slightly raised lateral ridges, apical part green on both sides, often deciduous. Sori 35 per pinna, linear-elliptic, ca. 1 mm, median on subtending vein; indusia grayish green, elliptic, membranous, entire and opening toward costa. Spores pale brown with lophate (costate) perispore, average exospore length 3136 ΅m.
* On rocks in forests; 1001000 m. Anhui, Fujian, Hunan, Jiangsu, Jiangxi, Yunnan, Zhejiang.
The type is from Jiangsu.
Asplenium kiangsuense is similar to a small form of A. normale, but differs in its scales with wider central band of occluded cells, the adaxially flat rachis, the green rachis tip, the smaller and more elliptic pinnae, and the perispore. The green rachis tip and the pinna shape make it also similar to A. adulterinum Milde, from which it differs in the blacker and wider zone of occluded cell lumina of its scales. Due to confusion with A. normale, the distribution of this taxon is not well known.
倒挂铁角蕨 dao gua tie jiao jue
Asplenium boreale (Ohwi ex Sa. Kurata) Nakaike; A. minus Blume; A. multijugum Wallich ex Mettenius; A. normale var. boreale Ohwi ex Sa. Kurata; A. normale var. shimurae H. Itτ; A. opacum Kunze; A. pavonicum Brackenridge; A. pseudonormale W. M. Chu & X.C. Zhang; A. shimurae (H. Itτ) Nakaike; A. trichomanes Linnaeus var. centrochinense Christ.
Plants 1540 cm tall. Rhizome erect, short, apex scaly; scales blackish brown, narrowly triangular, costate, with or without median opaque zone, margin fimbriate or entire. Fronds caespitose; stipe castaneous-brown to purplish black, shiny, 515(21) cm, terete to tri- or tetragonous; lamina linear-lanceolate, 1214(28) Χ 23(3.6) cm, 1-pinnate; pinnae 2030(44) pairs, alternate, sessile, middle pinnae trapeziform to oblong, 818 Χ 48 mm, base asymmetrical, acroscopic side truncate and subauriculate, basiscopic side narrowly cuneate, margin repand or crenate to serrate, apex obtuse; basal pinnae often somewhat reflexed, occasionally reduced. Costa with anadromously pinnate venation but with first basiscopic vein lacking, obscure, or faintly visible, veins simple or 2-forked. Fronds herbaceous to thinly papery, brownish green or grayish green when dry, (sub)glabrous; rachis castaneous-brown, subglabrous, often compressed after drying, adaxially with a deep furrow with rounded lateral edges, abaxially terete or keeled, often gemmiferous near apex. Sori 3 or 4(6) per pinna, linear-elliptic, 22.5(3) mm, median on subtending vein; indusium brown or grayish brown, semi-elliptic, membranous, entire, opening toward costa. Spores with lophate (cristate to alate) perispore, with perforated crests. Plants sexual: 2n = 72, 144, 216, or 288.
In soil or on rocks near streams, in forests; 6002500 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [India, Japan, Malaysia, Myanmar, Nepal, Philippines, Sri Lanka, Thailand, Vietnam; tropical Africa, Australia, Pacific islands].
The type is from Nepal.
In this Asplenium normale aggregate, three distinct taxa have been distinguished on the basis of the presence or absence of one to several buds on the rachis. At the tip of many fronds of typical A. normale sits a single gemma, often developed into a small plant. Plants without such buds have been described as A. normale var. boreale or A. boreale. Though the oldest name for non-gemmiferous plants would probably have to be A. minus (PRC!)[stet type info?]. The name A. shimurae, or A. normale var. shimurae, is used for plants with many gemmae per frond. Sterile hybrids between these taxa are known from Japan, and their flavonoid patterns are different. All three morphotypes occur in China (A. shimurae mainly in Yunnan), next to intermediate hybrids.
In this aggregate, diploid (China (Taiwan), S India, Malaysia, Nepal) and tetraploid chromosome numbers have been reported. Though Bir ([??Current Science 29: 445447]. 1960) and Nakaike ([??Bull. Nat. Sci. Mus. Ser. B 12: 3754]. 1986) mainly reported tetraploids for the E Himalaya, and Wang (1989) for China, we found tetraploid, hexaploid, and octoploid plants, next to intermediate hybrids.
Most of the typical gemmiferous plants of Asplenium normale are tetraploid, have scales with a central line of cells with more thickened and darker walls than the more marginal cells, have a deep furrow flanked by relatively broad and rounded edges at the adaxial side of the rachis, and lack the first basiscopic vein departing from the costa. Such tetraploid plants, as well as type material of A. multijugum from Nepal, have an average exospore length of 2630 ΅m. The absence of a basiscopic vein is a character A. normale shares with Hymenasplenium, in which two or more such veins are lacking.
Several morphologically similar plants have scales with the lumina of the central cells occluded and opaque (costate-opaque scales as in e.g., Asplenium trichomanes and A. kiangsuense) a shallow furrow or a flat rachis bordered by a narrow sharp rim (as in A. kiangsuense, not a wing as in A. trichomanes), and usually possess the first basiscopic vein on the costa. Sterile, morphologically intermediates between these plants and true A. normale are known from Guangdong. Because no clear correlations were found between the different ploidy levels (4x, 6x, 8x) and the morphotypes called A. normale, A. boreale, and A. shimurae, we refrain from recognizing them as species.
南海铁角蕨 nan hai tie jiao jue
Asplenium loriceum Christ; A. makinoi Hayata; A. sublongum Ching ex S. H. Wu.
Plants (30)5080 cm tall. Rhizome erect, short, apex scaly; scales reddish brown in center, narrowly triangular, costate with narrow pale denticulate margin. Fronds caespitose, rarely simple, usually pinnate with terminal pinna conforming to lateral ones; stipe 710 cm in simple fronds but 2025(40) cm in pinnate fronds, grayish green, scaly at base, subglabrous when old, adaxially grooved; lamina (in simple fronds) linear-lanceolate, 1535 Χ 1.52.5 cm, lamina in pinnate fronds ovate, 1835(50) Χ 1026 cm; pinnae 26 pairs, lower pinnae subopposite, with stalk 35(10) mm, lanceolate to linear-lanceolate, 1216(20) Χ 1.52.6 cm, base cuneate and decurrent on stalk, margin repand to sinuate, apex acuminate to caudate. Veins obvious, costa stramineous, notably raised on both sides, veins 2- or 3-forked, free. Fronds papery to subleathery when living, grayish green to brown when dry, abaxially with small, dark brown, fimbriate scales along costa, subglabrous when old; rachis grayish green, with small, dark brown, fimbriate scales. Sori linear, 510(15) mm, median to supra-median on acroscopic veinlets; indusium gray, linear, membranous, entire, opening toward costa, persistent.
Shaded places in dense forests or streamsides; 5002300 m. Guangdong, Hainan, Taiwan [Japan, Vietnam].
The type is from Taiwan.
Specimens of Asplenium formosae with sterile simple fronds are similar to A. scortechinii, but their fronds have an even to repand margin (notched in A. scortechinii) and a more prominent midrib.
兰屿铁角蕨 lan yu tie jiao jue
Asplenium wightianum Wallich ex Hooker var. microphyllum Matsumura & Hayata (1906), not Beddome (1864); Diplazium matsumurae (Christ) Kodama.
Plants 4080 cm tall. Rhizome erect. Fronds caespitose; stipe greenish stramineous, ca. 15 cm, base with subulate scales; lamina narrowly elliptic to ovate, ca. 35 Χ 12 cm, 1-pinnate, terminal apical segment similar to subapical pinnae or with 1 or 2 additional basal segments, ca. 4 cm; pinnae ca. 16 pairs, alternate, obliquely spreading, shortly stalked, lower pinnae not reduced, middle pinnae narrowly ovate-trullate, 710 Χ 11.3 cm, base asymmetrical, acroscopic side truncate to cuneate, basiscopic side narrowly cuneate, margin crenate-serrate, apex acuminate-caudate. Veins hardly visible, first basal basiscopic vein on costa lacking, suprabasal veins simple. Fronds herbaceous, grayish green when dry, subglabrous; rachis greenish stramineous. Sori linear, 34 mm, rarely diplazioid, medial; indusia linear, hyaline, entire, opening toward costa and persistent but not rolling back at maturity.
* On rocks in forests. Probably endemic to Taiwan.
Recently, Asplenium matsumurae has often been treated as a synonym of A. serricula Fιe, but that species has a more creeping rhizome, fewer pinnae, and a terminal pinna perfectly conforming to the lateral ones. Asplenium matsumurae is also superficially similar to members of the A. wrightii complex but it differs in its venation pattern with the first basal basiscopic vein lacking and more distal veins simple (not forked), its apex consisting of a subconforming pinna relatively similar to the subapical lateral pinnae, and its indusia not rolling back at maturity.
狭翅铁角蕨 xia chi tie jiao jue
Asplenium alatulum Ching; A. centrochinense Christ; A. centrochinense var. major Bonaparte; A. centrochinense f. maximum C. Christensen; A. duplicatoserratum Ching ex S. H. Wu; A. fujianense Ching (1981), not Ching ex S. H. Wu (1989); A. fujianense Ching ex S. H. Wu (1989), not Ching (1981); A. fujianensoides Viane; A. laui Ching; A. neomultijugum Ching [neomutijugum]; A. pseudowrightii Ching; A. serratissimum Ching ex S. H. Wu; A. subcrenatum Ching ex S. H. Wu; A. taiwanense Ching ex S. H. Wu; A. wrightii var. aristatoserrulatum Hayata; A. wrightii var. fauriei Christ; A. wrightioides Christ; Diplazium centrochinense (Christ) Tardieu.
Plants 3570(120) cm tall. Rhizome erect, short, scaly; scales brown, triangular to narrowly triangular, 57 Χ 11.2 mm, often costate with darker central zone and paler, denticulate glandular margin or long fibrillose. Fronds tufted; stipe dull to semi-shiny, grayish green to brown or stramineous-green, (12)2040(50) cm, with scales similar to those on rhizome, subglabrous with age; lamina narrowly ovate-trullate to elliptic, (20)4070(85) Χ (8)1525(30) cm, base truncate, apex acute, 1-pinnate; pinnae 1225 pairs, basal pinnae subopposite, others alternate, at an angle of ca. 50°60° to rachis, with stalks (2)48 mm, lower pinnae not much reduced, suprabasal pinnae narrowly ovate-trullate and often falcate, (5)1014(23) Χ (0.8)1.51.9(2.5) cm, base asymmetrical, acroscopic side truncate at an angle of (40°)55°75°(85°) to costa and often auriculate, basiscopic side cuneate, at an angle of (20°)30°40°(60°), becoming decurrent on rachis in apical part of lamina, margin variable, from almost entire to crenate-sinuate, denticulate-serrulate or serrate to dentate, more coarsely dentate toward pinna apex, apex acuminate. Veins (1 or)2-forked, with terminal hydathode. Fronds papery, green to brownish green when dry, subglabrous; rachis dull grayish brown to green or purple, with reddish brown, reduced and fimbriate scales or 2 or 3 cellular uniseriate gland-tipped hairs, or subglabrous, terete abaxially, winged toward apex. Sori linear, (3)912(14) mm, on acroscopic veinlets, medial to supramedial; indusia grayish brown to dark brown, linear, membranous, free margin entire but often glandular and rolling back at maturity, opening toward costa, persistent.
On rocks by streamsides in forests; 2001800 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [Japan, Korea, Vietnam].
The type is from Japan.
Most of the taxa put into synonymy of Asplenium matsumurae were based on variable characters, often of relatively small plants (nicely fitting a herbarium sheet). Within this aggregate, the name A. wrightii is generally applicable to the larger plants, but existing herbarium material is inadequate as it does not cover the morphological variation and cytology is usually unknown. A set of morphological and size characters separates this complex into four major, but probably artificial, groups. A first, including A. wrightii, has pinnae more than 10 cm, sori more than 8 mm and veins in the middle of the pinna 2-forked. A second group, including A. pseudowrightii, also has pinnae of more than 10 cm, but shorter sori (less than 8 mm), and veins only 1-forked in the middle of pinnae. Two groups with the largest pinnae less than 10 cm include A. wrightioides (with sori 611 mm) and A. centrochinense with sori less than 6 mm.
Japanese plants (Kurita, [??citation]. 1967; Mitsui, [??citation]. 1975) are octoploids (2n = 288), but Z. R. Wang ([??citation]. 1989) reported tetraploids (2n = 144) from Yunnan. Our recent flow-cytometry showed additional ploidy levels (10x and 12x), but because no correlation was found between ploidy levels and gross morphological characters, we refrain from recognizing separate species pending further micromorphological, cytological, and molecular research.
Asplenium pseudowrightii was erroneously ascribed to Hainan. This was due to a wrong stamp on the holotype sheet in PE, which is without a collection label. However, all isotypes have the same collectors label (numbered 37818), and collection labels state the plants origin: Guangdong: Xin Yi, Har Sick Joun, stream side, 15 Nov. 1934. Some plants from near the type locality are tetraploid (the lowest ploidy level in this complex), while others are 12x (highest ploidy level in this aggregate).
The type of Asplenium duplicatoserratum (S. S. Sin 469A) is also a syntype of A. centrochinense f. maximum. Asplenium duplicatoserratum is a large plant with serrate to biserrate pinnae similar to those of A. wrightii but with much shorter sori.
Asplenium Χwangii C. M. Kuo is a hybrid between A. wrightii s.l. and A. bullatum; it was described from Taiwan (Kuo, [??citation]. 1998) and may be present where these taxa co-occur (Fujian, Guizhou, Hunan, Sichuan, Taiwan, and Yunnan). Asplenium Χshikokianum Makino, the natural hexaploid hybrid between octoploid A. wrightii and tetraploid A. ritoense (described from Japan), is not uncommon where its parents grow together (i.e., Guizhou, Hunan, Sichuan, and Taiwan). Asplenium Χkenzoi Kurata is from the cross with A. prolongatum (Kurata, [??J. Geobot. 11: 68]. 1962).
网脉铁角蕨 wang mai tie jue
Asplenidictyum finlaysonianum (Wallich ex Hooker) J. Smith; Hemidictyum finlaysonianum (Wallich ex Hooker) T. Moore.
Plants 3050 cm tall. Rhizome erect, short, scaly; scales dark brown, narrowly triangular, entire. Fronds caespitose; stipe grayish green or stramineous, 1526 cm, with scales similar to those on rhizome, toward rachis with paleasters or subglabrous; lamina ovate-elliptic, 2032 Χ 1022 cm, 1-pinnate; pinnae 26(9) pairs, shortly stalked, lower pinnae not (much) reduced, ovate-lanceolate, 712 Χ 23.6 cm, base cuneate, gradually decurrent on stalk, margin entire to repand, apex acuminate and sometimes gemmiferous; terminal pinna similar to lateral ones but usually wider and more rhombic or hastate, 815 Χ 511 cm, its base often 3-lobed. Venation subflabellate with indistinct costa, veins faintly visible, almost parallel, ± anastomosing near margin. Frond papery or subleathery, brownish green when dry, with small scales along rachis and costa, becoming glabrous when old; rachis stramineous to grayish green, with sparse small narrowly triangular scales. Sori linear, variable in length, 545 mm, usually on acroscopic side of acroscopic veins; indusium yellowish to brown, linear, papery, entire, opening toward costa. Plants sexual polyploid: 2n = 144 or 288.
On wet rocks or tree trunks in dense forests; 7001100 m. Guangdong, Guangxi, Hainan, Xizang, S Yunnan [Bhutan, India, Indonesia, Malaysia, Myanmar, Nepal, Vietnam].
The syntypes are from Nepal and Kumaon.
Asplenium finlaysonianum is a Himalayan species with a typical morphology and venation pattern unlike any other taxon in the family. Though Bir ([??Current Science 29: 445447]. 1960) published a chromosome count showing this is a tetraploid (n = 72) in Sikkim, we found an octoploid plant in N Myanmar. The cytology of the Chinese plants is not known.
镰叶铁角蕨 lian ye tie jiao jue
Asplenium adiantoides (Linnaeus) C. Christensen (1905), not Lamarck (1786), nor Raddi (1819), nor Raoul (1844); A. falcatum Lamarck, nom. illeg.[check (Encycl. 2(1): 306. 1786)]; A. lofouense Christ; A. longjinense Ching & S. H. Wu; Tarachia falcata (Lamarck) C. Presl.
Plants 2060(90) cm tall. Rhizome shortly creeping to erect, scaly; scales dark brown, narrowly triangular with filiform apex, entire. Fronds caespitose; stipe dull or semi-shiny, grayish brown, 1035(45) cm, scaly at base, upward subglabrous, semiterete and adaxially grooved; lamina oblong to elliptical, 1035(45) Χ 816 cm, pinnate; pinnae (2)510(15) pairs, opposite or subopposite, distinctly stalked, lower pinnae not or only slightly reduced, median pinnae 410(15) Χ 1.22.2(2.8) cm, rhomboid to broadly lanceolate, ± falcate, base subsymmetrical, cuneate, acroscopic side sometimes subauriculate, basiscopic side narrowly cuneate, margin serrate often with long teeth, apex acuminate, terminal pinna conforming to lateral pinnae but often wider than subapical ones or with ascending pinnalike lobes at its base, 511 Χ 25(10) cm. Costa abaxially flat to slightly raised, adaxially with a longitudinal furrow (grooved). Veins prominent, several times forked, subflabellate, almost parallel, reaching margin. Frond (sub)leathery, dark brown when dry, pinna stalk and base with small dark brown scales becoming subglabrous when old; rachis dark gray-brown, semiterete and adaxially grooved. Sori linear, 13 cm, on acroscopic veins; indusia brown, linear, entire, opening toward costa. Plants sexual polyploid: 2n = 144, 216, or 360.
On rocks along streams or in forests; 300800 m. Guangdong, Guangxi, Guizhou, Hainan, Taiwan, SE Yunnan [India, Indonesia, Malaysia, Myanmar, Philippines, Sri Lanka, Vietnam; tropical Africa, Australia, Pacific islands (including New Zealand)].
The type is from New Zealand.
Asplenium polyodon is a widespread and variable aggregate species that needs more study before it can be split. Tetraploid chromosome numbers were reported from Australia, India (Darjeeling), Malaysia, New Zealand, and Sri Lanka. We found tetraploids and hexaploids in Uganda and China (Guangxi), and decaploids in Rιunion and China (Hainan). Plants with broad pinnae have once been identified as A. macrophyllum Swartz (in Schrader, J. Bot. 1800(2): 52. 1801), but the difference between this species and A. polyodon requires further study.
Chinese pinyin
Plants up to ca. 80 cm tall. Rhizome long creeping, scaly; scales dark brown, narrowly triangular, entire. Fronds separate along rhizome; stipe dull, dark grayish brown, 2040 cm, subglabrous, semiterete, grooved adaxially; lamina oblong to elliptical, 3045 Χ 1220 cm, pinnate; pinnae 1225(30) pairs, opposite or subopposite, distinctly stalked, lower pinnae not or only slightly reduced, median pinnae 713 Χ 0.81.5 cm, narrowly rhomboid-lanceolate, falcate, base asymmetrical, acroscopic side cuneate-truncate, basiscopic side narrowly cuneate, margin biserrate often with long teeth, apex long acuminate to caudate, terminal pinna reduced, apex pinnatifid. Costa abaxially slightly raised, adaxially with a shallow longitudinal furrow. Veins prominent, several times forked, distally in pinnae almost running parallel. Frond (sub)leathery, dark brown when dry, pinna stalk with small dark brown scales and becoming subglabrous when old; rachis dark gray-brown, semiterete, adaxially grooved. Sori linear, 812 mm, on acroscopic veins; indusia brown, linear, entire, opening toward costa. Perispore reticulate, without ridges.
On rocks in forests; ca. 1300 m. Hainan [Laos, Philippines, Vietnam].
The type is from the Philippines.
Asplenium lepturus is similar to A. polyodon but has more narrow and caudate pinnae, a widely creeping rhizome and a peculiar perispore pattern. It shares these characters with a number of tropical taxa, e.g., A. contiguum Kaulfuss from Hawaii, the neotropical A. serra Langsdorff & Fisher, the African A. friesiorum C. Christensen group, and A. nitens Swartz from Rιunion.
蒙自铁角蕨 men zi tie jue
Asplenium subtrapezoideum Ching ex S. H. Wu.
Plants 6070 cm tall. Rhizome suberect to creeping, with narrowly ovate-triangular, dark brown scales with narrow pale margin, up to 3 mm. Fronds approximate, subcaespitose; stipe dull grayish brown to stramineous, 1525(35) cm, with scales similar to those on rhizome; lamina narrowly triangular-ovate, 3540(45) Χ (10)1322 cm, 1-pinnate, apex triangular and pinnatifid, acute to acuminate; pinnae 1220 pairs, subopposite, shortly stalked (23 mm), basal pinnae only slightly reduced, largest pinnae (8)1014 Χ 1.53 cm, narrowly trapeziform-trullate and slightly falcate, base asymmetrical, acroscopic side truncate and often auriculate, basiscopic side narrowly cuneate, margin irregularly crenate-sinuate to crenate-serrate, apex acute-acuminate. Costa slightly sinuous, raised abaxially, adaxially with two ridges and sulcate; veins slender, 2- or 3-forked. Fronds firmly herbaceous, glaucous to yellowish brown-green or grayish green when dry, abaxially with minute, brown, hairlike scales; rachis grayish green to stramineous, with sparse scales, narrowly winged below apex, often gemmiferous below terminal segment. Sori linear, (0.8)12 cm, usually on acroscopic vein, at an angle of 10°25° with costa; indusia grayish brown, linear, membranous, margin with thin hyaline cells, entire, usually opening toward costa, rolling back and erose but persistent. Spores with average exospore length 3035 ΅m, perispore echinate.
In soil or on rocks in forests; 6001600 m. Guangxi, Hunan, Xizang, Yunnan [Vietnam].
Asplenium trapezoideum is a rare species, superficially similar to the more common A. wrightii, but it has a different rachis and costa structure, and is often proliferous near the apex. Asplenium trapezoideum is closely similar and shares its perispore pattern with the NE Indian A. khasianum Sledge, which may even be conspecific.
Asplenium subtrapezoideum, distinguished on size differences, is put into synonymy because fertile intermediate plants exist.
合生铁角蕨 he sheng tie jiao jue
Plants ca. 30 cm tall. Rhizome erect, short, scaly; scales dark brown to black, narrowly triangular, entire. Fronds caespitose; stipe dull, dark grayish brown, 1015 cm, with many dark brown, narrowly triangular to linear fimbriate scales; lamina linear, 1520 Χ ca. 3 cm, base truncate and slightly reduced, pinnatisect, apex acute; segments 1518 pairs, lower segments opposite, upper ones subopposite to alternate, middle pinnae 1.52 Χ 0.50.8 cm, narrowly ovate to elliptic, base adnate to rachis, upper segments connected by narrow wing, margin repand to sinuate, apex obtuse. Costa and veins obscure, veins simple or forked, reaching to margin. Fronds herbaceous, greenish brown when dry, abaxially with small hastate-fimbriate scales; rachis dark brown, with many narrowly triangular to linear fimbriate scales. Sori 3 or 4 per pinna, linear, 48 mm, approximately median on subtending vein; indusia linear, margin entire, opening toward costa, persistent.
* Guangdong (only known from the type).
Asplenium adnatum has scales similar to those of A. crinicaule but has adnate and obtuse pinnae. It is probably either a hybrid or an allopolyploid taxon derived from that species.
黑鳞铁角蕨 hei lin tie jiao jue
Plants ca. 30 cm tall. Rhizome erect, short, apex scaly; scales blackish brown, narrowly triangular, with broad, fimbriate and hastate base, apex filiform. Fronds caespitose; stipe castaneous to dark brown, 57 cm, with scales similar to those on rhizome; lamina linear, 2023 Χ 23 cm, attenuate to both ends, 1-pinnate; pinnae ca. 25 pairs, basal ones (sub)opposite, upper ones alternate, sessile, middle pinnae oblong-elliptic, 11.5 Χ ca. 0.4 cm, base asymmetrical, acroscopic side truncate, basiscopic side cuneate, margin crenate-serrate, apex truncate-obtuse; lower pinnae gradually reduced. Venation anadromously pinnate, obscure, veins simple or forked. Fronds herbaceous-leathery, brownish green when dry, both surfaces with hastate-stellate scales; rachis dark purplish brown, semiterete, adaxially sulcate, ridges rounded and without wings, with hastate-stellate scales. Sori 68(or 9) per pinna, linear, 34 mm, on acroscopic veinlets; indusia yellowish brown, broadly linear, membranous, entire, opening toward costa.
* On rocks in forests; ca. 1000 m. Guizhou (Duyun, Tonren).
Asplenium asterolepis is similar, and perhaps conspecific, to A. crinicaule but has smaller, more oblong pinnae. Its scales are deltoid-tridentate with a very long central apical tail, as those of A. crinicaule. At present, this taxon seems to be known only from the type collection and may represent just a local variant of A. crinicaule.
毛轴铁角蕨 mao zhou tie jiao jue
Asplenium beddomei Mettenius ex Kuhn; A. hancei Baker; A. polytrichum Christ; A. saigonense Matthew & Christ.
Plants 2040 cm tall. Rhizome erect, short, scaly; scales blackish brown, narrowly triangular to deltoid-tridentate with long apical tail. Fronds caespitose; stipe grayish brown to purplish black, 512 cm, semiterete, adaxially grooved, with blackish brown deltoid-tridentate scales with long apical tail; lamina lanceolate to linear-lanceolate, 1030 Χ 3.58 cm, gradually reduced at base, apex acute, 1-pinnate; pinnae 1830(40) pairs, alternate or lower ones (sub)opposite, almost sessile to shortly stalked; basal pinnae slightly reduced, deltoid-triangular to ovate, apex obtuse; middle pinnae narrowly subtriangular to rhomboid-elliptic, 24(5.5) Χ 0.81.5 cm, base asymmetrical, acroscopic side truncate and auriculate, basiscopic side narrowly cuneate, margin irregularly biserrate-crenate, apex acute. Veins obvious, mostly 2-forked, 3-forked, or simple, not reaching margin. Fronds papery, dark brown when dry, both surfaces with dark brown stellate or hairlike small scales, subglabrous when old; rachis grayish brown, semiterete, adaxially grooved, with rigid hairlike scales, rarely with subapical gemma. Sori linear to crescent-shaped, 48 mm, on acroscopic veinlets; indusia yellowish brown to grayish brown, linear, membranous, entire, opening toward costa or to first acroscopic vein, often rolling back at maturity, persistent. Perispore costate, with average exospore length 4044 (dodecaploids). Plants sexual polyploid: 2n = 144, 288, or 432.
On rocks at streamsides, in forests; 1003000 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Jiangxi, Sichuan, Xizang, Yunnan [India, Malaysia, Myanmar, Philippines, Thailand, Vietnam; Australia].
The type is from Guangdong (Dinghu Shan).
Tetraploids plants (n = 72) of Asplenium crinicaule were reported from India (Bir, [??Current Science 29: 445447]. 1960), but in China, this is an aggregate of octo- and dodecaploid taxa. At present, octoploids are known from S India (Kuriachan, [citation?]. 1968), Sichuan (Wang, [??in Shing, K. H., Kramer, K. U. Proc. Int. Symp. Syst. Pterid. 133134]. 1989), Guangdong, and Guizhou, and dodecaploids from Guizhou and Guangdong.
Plants are variable in frond and pinna shape, even at the type locality at Dinghu Shan, where the species is dodecaploid. Members of this aggregate are similar to Asplenium pellucidum Lamarck, which is octoploid in Rιunion, but has more narrow pinnae with an acuminate, apex and many strongly reduced basal pinnae. Asplenium horridum Kaulfuss has similar scales. We prefer to treat this taxon as an aggregate species until the ploidy of plants at the different type localities is better known.
胎生铁角蕨 tai sheng tie jiao jue
Asplenium planicaule Wallich ex Mettenius (1859), not E. J. Lowe (1858); A. laciniatum D. Don var. planicaule Clarke[IPNI: C. Christensen]; A. wuyishanicum Ching; A. yoshinagae Makino subsp. indicum (Sledge) Fraser-Jenkins; A. yoshinagae var. indicum (Sledge) Ching & S. K. Wu; A. yoshinagae var. planicaule (Clarke[C. Christensen?]) C. V. Morton.
Plants 1025 cm tall. Rhizome erect, short, scaly; scales dark to reddish brown, narrowly triangular, entire, apex acuminate. Fronds caespitose; stipe grayish brown or grayish green to stramineous, 510 cm, adaxially grooved longitudinally, with small scales similar to rhizome scales, subglabrous when old; lamina narrowly triangular, 725 Χ 25 cm, apex acuminate, 1-pinnate; pinnae 820 pairs, alternate or lower ones opposite, shortly stalked, basal pinnae not or slightly reduced, middle pinnae 12.5 Χ 0.51.3 cm, rhomboid to trapezoid or subdimidiate, ± falcate, base asymmetrical, first acroscopic lobe larger than first basiscopic lobe, acroscopic side truncate, almost parallel to rachis and auriculate, basiscopic side narrowly cuneate (1/41/3 cut off) to almost parallel to costa, margin irregularly bicrenate, apex acute to acuminate, rarely obtuse. Veins obvious, grooved adaxially, basal acroscopic vein multi-forked, other veins 1- or 2-forked. Fronds subleathery, grass-green to stramineous when dry, adaxially with wrinkles above veins, with small, brown, narrowly triangular and apically filiform scales on abaxial surface and pinna stalks, subglabrous when old; rachis (when dry) abaxially grayish castaneous to green-stramineous for most of its length, and with small narrowly triangular and apically filiform scales, adaxially longitudinally valleculate, often with scaly gemmae or juvenile plants on pinna stalks or rachis. Sori linear, 37 mm, from near costa running nearly to margin, distal sori close to rachis; indusia grayish brown, linear, entire, opening toward costa or first acroscopic vein. Spores with lophate (costate) perispore, average exospore length 3136 ΅m. Plants tetraploid: 2n = 144.
On wet rocks or tree trunks in forests; 6002700 m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hunan, Jiangxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Bhutan, India, Myanmar, Nepal, Philippines, Sri Lanka, Thailand, Vietnam].
The type is from Nepal.
Asplenium indicum is a sexual tetraploid plant in India, the Himalaya (Mehra & Bir, [??citation]. 1957; Bir, [??Current Science 29: 445447]. 1960; [??Curr. Sci. 31: 248250. 1962; Caryologia 18: 107115. 1965]), and China (Xizang, Guizhou). It is morphologically very variable, and similar to A. yoshinagae with which it is often united to form an aggregate taxon. Because they are genetically isolated and form sterile hybrids when growing together, we treat these taxa as separate species. In China, A. indicum is usually a smaller plant, with smaller spores and stomates than A. yoshinagae, from which it can be distinguished by a chromosome count or flow-cytometry. The distinction based on the presence of gemmae is highly unreliable, at least in China, where all tetraploids checked were gemmiferous. The gemmae are formed on the pinnae stalks, close to the rachis, rarely on the rachis itself. Another very similar taxon, A. gueinzianum, is also gemmiferous, but produces buds on the surface of the pinnae. Due to confusion among these taxa, the exact distribution of this species is not well documented.
棕鳞铁角蕨 zon lin tie jiao jue
Asplenium indicum var. yoshinagae (Makino) Ching & Wu; A. planicaule E. J. Lowe var. yoshinagae (Makino) Tagawa; A. tibeticum Ching; Tarachia yoshinagae (Makino) H. Itτ.
Plants 2045 cm tall. Rhizome erect, short, scaly; scales dark to reddish brown, narrowly triangular, entire, apex acuminate. Fronds caespitose; stipe grayish brown or grayish green to stramineous, 1020 cm, adaxially grooved longitudinally, with small scales similar to rhizome scales, subglabrous when old; lamina narrowly triangular, 1230 Χ 47 cm, apex acuminate, 1-pinnate; pinnae 820 pairs, alternate or lower ones opposite, shortly stalked, basal pinnae not or slightly reduced, middle pinnae 23.5 Χ 11.3 cm, rhomboid to trapezoid or subdimidiate, ± falcate, base asymmetrical, first acroscopic lobe much larger than first basiscopic lobe, acroscopic side truncate, almost parallel to rachis and auriculate, basiscopic side narrowly cuneate (1/41/3 cut off) to almost parallel to costa, margin irregularly bicrenate, apex acute to acuminate, rarely obtuse. Veins obvious, grooved adaxially, basal acroscopic vein multi-forked, other veins 1- or 2-forked. Fronds subleathery, grass-green to stramineous when dry, adaxially with wrinkles above veins, small, brown, narrowly triangular and apically filiform scales on abaxial surface and pinna stalks, subglabrous when old; rachis (when dry) abaxially grayish castaneous to green-stramineous for most of its length, and with small narrowly triangular and apically filiform scales, adaxially longitudinally valleculate, gemmae or juvenile plants near rachis on pinna stalks. Sori linear, 48 mm, from near costa running nearly to margin, distal sori close to rachis; indusia grayish brown, linear, entire, opening toward costa or first acroscopic vein. Spores with lophate (costate) perispore, average exospore length 3640 ΅m. Plants sexual octoploid: 2n = 288.
On wet rocks or tree trunks; 6002500 m. Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Sichuan, Xizang, Yunnan, Zhejiang [India, Japan, Vietnam].
The type is from Japan.
Asplenium yoshinagae is a sexual octoploid plant in S India (Kuriachan, [??citation]. 1965), Japan (Shimura & Takiguchi, [??citation]. 1979), and China (Guangdong, Guangxi, Xizang, and Yunnan). It is morphologically very variable, and similar to A. indicum (see above). In China, A. yoshinagae is usually a larger and more vigorous plant with larger spores and stomates than A. indicum. They can easily be distinguished by a chromosome count or by flow-cytometry. The presence of gemmae is a highly unreliable character, at least in China, where most octoploids checked were not gemmiferous. Due to confusion among these taxa, the exact distribution of this species is not well documented.
Asplenium tibeticum, only known from the type collection from Xizang, has leathery fronds morphologically almost intermediate between A. yoshinagae and A. gueinzianum; however, its long slender stipe scales, and epidermal and soral cell patterns agree with those of A. yoshinagae, and its average exospore length of 40 ΅m also fits for this taxon.
撕裂铁角蕨 si lie tie jie jue
Plants 2535 cm tall. Rhizome erect, short, apex scaly; scales brown or reddish brown, narrowly triangular, entire. Fronds caespitose; stipe dull gray-brown, 46 cm, with many narrowly triangular, costate scales with dark brown central zone and almost hyaline margin, adaxially sulcate; lamina narrowly triangular to linear, 2030 Χ 34 cm, apex acuminate, 1-pinnate to pinnate-pinnatifid; pinnae 2026 pairs, alternate or subopposite, shortly stalked, middle pinnae obliquely rhomboid, 1.42 Χ 0.60.9 cm, apex acute, base asymmetrical, acroscopic side truncate and auriculate-cordate, basiscopic side narrowly cuneate, margins irregularly and deeply bisinuate-bicrenate, basal acroscopic segment most developed, flabellate-obdeltoid, often almost free from rest, first basiscopic segment below second acroscopic lobe; basal pinnae gradually and slightly reduced, often deflexed. Veins visible, basal 1/32/3 of costa forming lower margin, lateral veins 2-forked. Fronds herbaceous, green when dry, glabrous adaxially but usually with a small gemma near apex, abaxially with brown small scales or subglabrous; rachis gray-brown to stramineous, with scales similar to those of stipe, subglabrous when old, adaxially sulcate. Sori 58 per pinna, linear, 23.5 mm, median on acroscopic veinlets; indusia yellowish green to grayish brown, oval-linear, entire, mostly opening toward costa. Spores with lophate (cristate-alate) perispore. Plants sexual: 2n = 144.
On wet rocks at streamsides; 15002600 m. Taiwan, Xizang, Yunnan [Bhutan, N India, N Myanmar, Nepal, Vietnam].
The type is from Nepal.
The name Asplenium laciniatum D. Don (Prodr. Fl. Nepal. 8. 1825; Tarachia laciniatum (D. Don) C. Presl), which has been widely and persistently used for this taxon, does not include its type and should thus be rejected (Vienna Code, Art. 57.1). The problem with the application of the name was caused by Hooker ([??Sp. Filic. 3:??]. 1860), who was followed by most later authors. This was first exposed by Morton ([citation?]. 1973) and was discussed by Sleep and Reichstein ([citation?]. 1984) and Viane and Reichstein ([citation?]. 1986) in relation to the typification of A. varians.
Fronds of Asplenium gueinzianum are similar to those of A. indicum and A. yoshinagae, as well as to those of the A. varians complex, but have costate scales with hyaline borders and small buds or plantlets on the frond surface near the pinna apex (not on the rachis as in A. indicum/yoshinagae).
西南铁角蕨 xi nan tie jiao jue
Trichomanes aethiopicum N. L. Burman, Fl. Ind. (Prodr. Fl. Cap.) 28. 1768; Asplenium adiantoides Lamarck; A. denticulatum Blume; A. furcatum Thunberg; A. praemorsum Swartz; Tarachia furcata (Thunberg) C. Presl.
Plants 2545 cm tall. Rhizome short, erect to shortly creeping, scaly; scales dark reddish brown to black, narrowly triangular with a long filiform apical tail. Fronds caespitose; stipe 515(22) cm, adaxially grayish green and sulcate, abaxially dark brown to black, with many reddish to dark brown, narrowly triangular scales with a long filiform apex; lamina narrowly ovate to narrowly triangular, 1028 Χ 4.58 cm, apex acuminate, pinnate-pinnatifid to 2-pinnate; pinnae 1015 pairs, subopposite or alternate, lower 1 or 2 pairs slightly reduced, subsessile, trullate-rhomboid with 1 or 2 (almost) free pinnules at base, 2.54.5 Χ 12.5 cm, base asymmetrical, broadly cuneate, apex acuminate; pinnules 24 pairs, obtrullate-elliptic or subflabellate, 412 Χ 25 mm, margin dentate, lateral sides entire, apex obtuse. Venation subflabellate, obvious abaxially, veins slender and forked, not reaching margin, costa grooved adaxially. Fronds leathery, adaxially dark green when dry and often grooved, abaxially brownish green, with reddish brown, fibrillose scales, subglabrous when old; rachis green but often becoming dark toward base abaxially, adaxially sulcate and densely scaly with fibrillose scales. Sori 25 per segment, linear, 38 mm, median on subtending veins; indusia grayish yellow, linear, membranous, entire, opening toward major veins or to costa, persistent. Plants polyploid, sexual or agamosporous.
On rocks under mixed forests; 10002600 m. Hunan, Jiangxi, Sichuan, Yunnan [India, Indonesia, Malaysia, Myanmar, Philippines, Thailand, Vietnam; tropical Africa, tropical America, Australia, Pacific islands (Macronesia, Polynesia: Hawaii)].
Asplenium aethiopicum is a morphologically, ecologically, and karyologically very variable aggregate for which we prefer to use the oldest available epithet. Braithwaite ([??citation]. 1964; [??Bot. J. Linn. Soc. 93: 348378]. 1986), Panigrahi ([??Phytologia 31: ??]. 1975), Manton et al. ([??Bull. Brit. Mus. (Nat. Hist.), Bot. Ser. 15: 123161]. 1986), and Ormonde ([??Acta Bot. Malacitana 16: 293315]. 1991) have shown that A. aethiopicum s.l. is an aggregate of at least three sexual (A. aethiopicum subsp. tripinnatum (Baker) A. F. Braithwaite (4x), A. aethiopicum subsp. aethiopicum (8x), and A. aethiopicum subsp. dodecaploideum A. F. Braithwaite (12x)) and up to five apomictic taxa. Most of these cytotypes are common in Africa, but sexual tetraploids and dodecaploids are also known from Asia and America, where Swartzs name, A. praemorsum, is used for the aggregate. Besides differing in chromosome number and pairing behavior, the members of this aggregate can also be separated on the basis of their exospore size. The agamosporous taxa can be distinguished from the sexual by the length/width ratio of their spores: agamosporous taxa have subglobose spores with a ratio smaller than 1.4, sexual taxa have reniform to plano-convex spores with a ratio larger than 1.4. Based on Braithwaites ([??citation]. 1964) key, Chinese specimens are sexual and belong to A. aethiopicum subsp. dodecaploideum. We found hybrids with A. austrochinense in Jiangxi.
斜裂铁角蕨 xie lie tie jiao jue
Plants 4045 cm tall. Rhizome erect, short, scaly; scales brown, narrowly triangular, margins dentate to fimbriate. Fronds caespitose; stipe grayish brown, 2035 cm, sulcate adaxially, when young with dark brown, narrowly triangular to ovate scales with several rows of marginal cells less thickened than in central zone, subglabrous when old; lamina ovate-elliptic, 2030 Χ 68 cm, apex acute, pinnate-pinnatifid; pinnae 1015 pairs, with stalks 24 mm, not much reduced toward base, narrowly triangular to rhomboid, 33.5 Χ 12.2 cm, base asymmetrical, broadly cuneate, margin dentate, apex acuminate; segments 3 or 4 pairs, rectangular to elliptic, 611 Χ 38 mm, basal acroscopic segment largest, flabellate, apex obtuse and dentate, lateral sides entire. Veins obvious on both sides, costa shallowly sulcate adaxially, veins 2- or 3-forked, running almost to margin. Fronds herbaceous to papery, brownish green when dry, with supravascular ridges on lamina adaxially, rachis greenish gray, sulcate adaxially, abaxially grayish brown, with sparse dark brown, narrowly triangular scales. Sori 24 per segment, linear, 2.55 mm, median on subtending veins; indusia gray-brown, linear, entire, opening toward major veins or costa, persistent. Plants sexual tetraploid: 2n = 144*.
* On rocky cliffs in forests; 5001000 m. Hainan.
Asplenium pseudopraemorsum is similar to A. aethiopicum in general aspect and outline, but lacks the reddish brown fimbriate scales and has shorter sori. Asplenium protractum Tardieu & Ching is also similar but has more subleathery fronds with shorter sori.
石生铁角蕨 shi sheng tie jiao jue
Asplenium affine Swartz var. sinense Christ; A. comptum Hance (1866), not Kunze ex T. Moore & Houlston (1852); A. dimidiatum Swartz var. comptum Baker.
Plants 2050 cm tall. Rhizome erect, short, scaly; scales dark brown, narrowly triangular, margins dentate-fimbriate. Fronds caespitose; stipe grayish green to black, 1022 cm, base scaly, more sparsely so upward, adaxially sulcate; lamina triangular-ovate to narrowly triangular, 1228 Χ 511 cm, pinnate or pinnate-pinnatifid to 2-pinnate at base and gradually becoming pinnate toward apex, apex acute; terminal apical pinna simple or pinnatifid; pinnae 512(14) pairs, simple or pinnate (basal pinnae), opposite or alternate, stalk 512 mm; basal pinnae usually largest, 36 Χ 23 cm, rhomboid or triangular to narrowly triangular, base asymmetrical, acroscopic side truncate to cuneate, basiscopic side cuneate, margin irregularly crenate-sinuate, apex acute to acuminate; segments 13, rhomboid to elliptic or obovate, 0.62 Χ 0.31.2 cm, basal acroscopic segment largest. Costa shallowly sulcate adaxially, lateral veins flabellately forking, not reaching margin. Fronds leathery, dark brown when dry; rachis dark brown to grayish black at base, becoming green toward apex, with dark brown, narrowly triangular scales with lateral fimbriae and filiform apex, subglabrous when old, adaxially sulcate. Sori 36 per segment (812 on basal acroscopic segment), linear, 415 mm; indusia brown, linear, thickly membranous, entire, opening toward costa or major veins. Spores with dark brown, lophate perispore, average exospore length 2629 ΅m (in tetraploids) or 4045 ΅m (in octoploids).
On limestone rocks in forests; 3001300 m. Guangdong, Guangxi, Guizhou, S Hunan, Sichuan, Yunnan [Vietnam].
The type is from Guizhou.
Asplenium saxicola is an aggregate of tetraploid and octoploid plants with fronds similar, though more divided at the base, to those of A. dimidiatum Swartz (America) and A. hemitomum Hieronymus and A. megalura Hieronymus (Africa). The scales are similar to those of A. aethiopicum but have thicker cell walls and a shorter apical tail. The ploidy level of plants from the type locality is not known. Tetraploids were found in Guangxi, and octoploids in Guangxi and in Yunnan.
东南铁角蕨 dong nan tie jiao jue
Asplenium formosanum Baker; A. hancockii Baker (1885), not Maximowicz (1883); Tarachia austrochinensis sensu B. Itτ[(Ching) H. Itτ per IPNI; should be a synonym of sp. 52; though FRPS (4(2): 68. 1999) says non Asplenium austrochinense Ching, 1931].
Plants 1520 cm tall. Rhizome erect, short, scaly; scales dark brown, narrowly triangular, subentire. Fronds caespitose; stipe green or base brownish gray, 410 cm, adaxially sulcate, with small reddish brown narrowly triangular scales or subglabrous; lamina ovate-triangular, 610 Χ 34 cm, apex acute, pinnate-pinnatisect to almost bipinnate; pinnae 59 pairs, subopposite or alternate, stalk 23 mm, basal pinnae not reduced, rhombic, 1.83 Χ 12 cm, base slightly asymmetrical, broadly cuneate, apex subacute; segments 1 or 2 pairs, elliptic, 311 Χ 2.55 mm, basal acroscopic lobe largest, base decurrent on costa, crenate to sinuate, apex obtuse to truncate. Costa shallowly grooved adaxially, veins anadromously forking. Fronds leathery, dark green when dry; rachis grayish stramineous or green, sulcate adaxially, with dark brown, narrowly triangular scales, subglabrous when old. Sori 14 per segment, median to distal on subtending veins, not reaching margin, oval-linear, 35 mm; indusia brown, oval-linear, membranous, entire, opening toward costa, persistent.
* On wet rocks in forests; ca. 700 m. Anhui, Fujian, Jiangxi, Taiwan, Zhejiang.
The type is from Taiwan.
The relationship of Asplenium oldhamii to A. austrochinense requires further study.
匙形铁角蕨 chi xing tie jiao jue
Asplenium spathulinum J. Smith ex Hooker (1860), not Kunze (1848).
Plants 3065 cm tall. Rhizome erect or ascending, scaly; scales brown, narrowly triangular, margins subdentate. Fronds caespitose; stipe dull grayish brown to gray-green, semiterete, 1330 cm, subglabrous, adaxially sulcate; lamina triangular to ovate, 2136 Χ 1020 cm, not much reduced at base, apex acute-acuminate, bipinnate; pinnae 1218 pairs, subopposite to alternate, stalks 23 mm, basal pinnae largest or slightly reduced, narrowly triangular, 714 Χ 3.54.5 cm, apex variable, from acute to caudate, 1-pinnate; pinnules 48 pairs, anadromous, free to subsessile, from ovate-elliptic to rhombic or cuneiform, apex obtuse, basal acroscopic pinnule about equal to basal basiscopic pinnule, basal pair largest, 1.53 Χ 11.3 cm, base subsymmetrical, cuneate, distal segments decurrent on rachis, lateral sides entire, outer margin dentate to crenate. Costa obvious on both sides, raised abaxially, adaxially shallowly grooved with raised supravascular ridge, veins 2- or 3-forked and not reaching margin. Fronds herbaceous, brownish green when dry, subglabrous; rachis grayish brown to gray-green, adaxially sulcate, apical part if rachis often gemmiferous. Sori 14 per pinnule, not reaching margin, linear, 510 mm; indusia brownish, linear, membranous, entire, opening toward costa or costule, persistent.
Epiphytic in forests; 6001400 m. Hainan [Indonesia, Malaysia, Philippines, Sri Lanka; Mascarene Islands].
The type is from Mauritius.
Hookers name, Asplenium spathulinum, has often been used to describe this variable taxon (e.g., Beddome, [??citation]. 1864; Tardieu & Ching, [??citation]. 1936; Chun et al. [??citation]. 1964; Copeland, [??citation]. 1960; Holttum, [??citation]. 1955; [??citation]. 1968; S. H. Wu, [??citation]. 1999; T. L. Wu et al., [??citation]. 2006). Manton & Sledge ([??citation]. 1954) showed that A. affine is an aggregate of an octoploid and a dodecaploid species (S India and Sri Lanka). Plants from closer to the type locality, in Rιunion, were dodecaploid. Because specimens from the Hainan populations have the same ploidy level (12x), we use the name A. affine following Sledges ([??citation]. 1965) concept, based on morphological and cytological study. Mature A. affine plants can be very similar to juvenile specimens of A. pseudolaserpitiifolium and its relatives.
烏来铁角蕨 wu lai tie jiao jue
Asplenium arisanense Tagawa; Tarachia cuneatiformis (Christ) H. Itτ.
Plants 5060 cm tall. Rhizome ascending or shortly creeping, scaly; scales brown to black, narrowly triangular, entire. Fronds subcaespitose; stipe dull, dark purplish to gray-brown, semiterete, 1924 cm, adaxially sulcate, base scaly, upward subglabrous; lamina narrowly triangular to ovate, 1530(40) Χ 58 cm, bipinnate at base, apex acuminate; pinnae 1420 pairs, opposite to alternate, stalks ca. 2 mm, basal pinnae not reduced, narrowly triangular to ovate, 56 Χ ca. 2 cm, apex obtuse to subacute, serrate; basal pinnae 1-pinnate, free pinnules 13(or 4) pairs, basal acroscopic pinnule largest, ovate-trullate, 1014 Χ 56 mm, apex obtuse or truncate, base cuneate, adnate to costa toward apex, apical pinnae pinnatifid or simple, acroscopic side auriculate. Costa flat to slightly raised adaxially. Veins obvious on both surfaces, anadromous, 2- or 3-forked, not reaching margin. Fronds papery, brown when dry, lamina subglabrous; rachis semiterete, dull, dark purplish to gray-brown abaxially, green toward apex, with small subhastate to stellate reduced scales or hairs to subglabrous, adaxially sulcate and often gemmiferous near apex. Sori 13 on basal pinnule, median on subtending vein, not reaching margin, linear, 59 mm; indusia brown to dark brown, linear, membranous, entire, opening toward costa or costule, persistent.
* On wet rocks or epiphytic; 500-700 m. Taiwan.
Asplenium cuneatiforme is similar to A. affine, which has a wider and more divided frond, and to the Pacific A. lobulatum Mettenius ex Kuhn with a creeping rhizome and a less divided lamina.
瑞丽铁角蕨 rui li tie jiao jue
Asplenium laciniatum D. Don var. crinigerum Beddome.
Plants up to 20 cm tall. Rhizome short, erect, or ascending; scales dark brown to black, narrowly triangular, entire or subfimbriate. Stipe stramineous, 24 cm, semiterete to terete, with spreading, narrowly triangular, dark brown scales, with long acuminate apex. Lamina narrowly elliptic to lanceolate, 1218 Χ 23.5 cm, base reduced, 2-pinnate to 3-pinnatifid, apex acute; pinnae 1520 pairs, subopposite to alternate, stalk ca. 1 mm, lower pinnae reduced, middle pinnae elliptic-subtriangular, 1.52.3 Χ 0.91.2 cm, base asymmetrical, acroscopic side truncate, basiscopic side cuneate, apex obtuse to subacute, 1-pinnate or pinnatisect; pinnules 3 pairs, anadromous, basal acroscopic pinnule largest, cuneate-spatulate, 68 Χ 34 mm, base cuneate and decurrent on costa wing, apex obtuse-truncate and serrate-lacerate, with 24 teeth up to 7 Χ 3.5 mm. Fronds subleathery, green, edges often revolute when dry, with narrow scales on veins, lamina subglabrous; rachis stramineous or green, sulcate adaxially, with dark brown, narrowly triangular, caudate scales. Costa minutely scaly abaxially, sulcate but with raised supravascular ridge, veins anadromous, not reaching margin. Sori 1 or 2(or 3) per pinnule, median on subtending veinlets, elliptic, 24 mm; indusia gray, elliptic, membranous, entire, opening toward costa or costule.
On rocks. Yunnan [Bhutan, NE and S India, N Myanmar, N Thailand].
The type is from Thailand.
Asplenium rockii is a distinct but rare species, which according to flow cytometric analysis, is hexaploid in China.
华南铁角蕨 hua nan tie jiao jue
Asplenium consimile Ching ex S. H. Wu (1989), not Rιmy (1854); A. jiulungense Ching; A. nanchuanense Ching & Z. Y. Liu; A. pseudowilfordii Tagawa; A. wilfordii Mettenius ex Kuhn var. austrochinense (Ching) Tagawa.
Plants 3040 cm tall. Rhizome erect, short, apex scaly; scales dark to medium brown, narrowly triangular, fimbriate in basal part to subentire near apex. Fronds caespitose; stipe up to 20 cm, semiterete, base abaxially dull, dark purplish to gray-yellow when dry, adaxially green and sulcate, base with scales similar to those on rhizome, toward rachis with gradually smaller and more hastate-stellate scales with fimbriae; lamina triangular-ovate to narrowly triangular, 1826 Χ 610 cm, base cuneate to truncate, apex acute-acuminate, 2-pinnatifid to 3-pinnatifid; pinnae 1015 pairs, subopposite to alternate, stalk 34 mm, basal pinnae not reduced, narrowly triangular-ovate, 4.58 Χ 1.73 cm, apex acute to caudate, pinnate to 2-pinnatifid; pinnules 35 pairs, anadromous, basal acroscopic pinnule largest, free or adnate to costa, rhombic, 12 Χ 0.61.2 cm, apex obtuse-truncate to subacute, basal pinnule simple to pinnatifid and with 2 or 3 segments, base cuneate or decurrent on costa in upper pinnae, margin serrate-crenate to sinuate, teeth usually obtuse, costa sulcate adaxially, narrowly winged. Veins obvious abaxially, obscure but occasionally raised adaxially, venation anadromous, veins subflabellate, not reaching margin. Fronds herbaceous to leathery, green, or yellowish brown when dry, lamina with reduced, hastate-stellate, and fimbriate scales or subglabrous; rachis semiterete, green, or more rarely purplish brown or gray, sulcate and green adaxially. Sori 26(9) per pinnule, short, linear, 35 mm; indusia pale brown, linear, firm, membranous, entire, opening toward costule or costa, persistent. Plants octoploid: 2n = 288.
On wet rocks or moss cushions on trees in dense forests; 4001100 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan, SE Yunnan, Zhejiang [Japan, Vietnam].
The type is from Guangdong.
Plants of Asplenium austrochinense from Guangdong are octoploid like the Japanese (Shimura & Takiguchi, [??citation]. 1979), and scales are similar in Chinese and Japanese specimens. Upon drought stress, the margins of the frond tend to roll inward, protecting the abaxial epidermis with the stomates. The name Asplenium consimile Ching was used to describe plants with a somewhat thinner lamina.
海南铁角蕨 hai nan tie jiao jue
Plants 2440 cm tall. Rhizome erect, short, apex scaly; scales dark brown, narrowly triangular, entire. Fronds caespitose; stipe gray to dark brown, 410(15) cm, adaxially sulcate, with brown narrowly triangular or stellate scales, often becoming subglabrous; lamina narrowly triangular to ovate, 1326 Χ 38 cm, apex acuminate-caudate, bipinnate; pinnae 1016 pairs, subopposite to alternate, stalk up to ca. 2 mm, basal pinnae largest or slightly reduced, middle pinnae ovate, 2.54(7) Χ 1.21.6 cm, apex obtuse to subacute, 1-pinnate; pinnules 24(7) pairs, anadromous, first acroscopic pinnule largest, narrowly obovate-obtrullate, 810 Χ 46 mm, distal pinnules narrowly cuneiform, basiscopic pinnules cuneiform-falcate, base narrowly cuneate, almost sessile, lateral side entire, apex serrate and obtuse. Costa sulcate with raised supravascular ridge adaxially, green. Veins distinct, raised adaxially, anadromously branching, not reaching margin. Fronds firmly herbaceous, brownish green when dry; rachis gray to dark brown abaxially, becoming green toward apex, subglabrous, adaxially sulcate. Sori 25 per pinnule, linear, 26 mm; indusia pale brown, linear, membranous, entire, opening toward costule, persistent.
On wet rocks at streamsides in forests; 400700 m. Hainan [Thailand, Vietnam].
The type is from Hainan.
Asplenium hainanense is similar to A. gracilifolium Copeland and to small forms of A. affine, but never proliferous.
俅铁角蕨 qiu jiang jiao jue
Replaced synonym: Asplenium qiujiangense Ching ex S. H. Wu, Bull. Bot. Res., Harbin 9(2): 90. 1989, not Asplenium qiujiangense (Ching & Fu) Nakaike, Bull. Natl. Sci. Mus., Tokyo B, 12(2): 45. 1986; A. dulongjiangense Y. F. Deng (2003), not Viane (1991).
Plants 50100 cm tall. Rhizome shortly creeping or erect, scaly; scales purplish brown, narrowly triangular, entire. Fronds caespitose; stipe 3038 cm, semiterete, base scaly, upward subglabrous, dull, grayish green, adaxially sulcate; lamina ovate-triangular, 4560 Χ 2630 cm, apex acute-acuminate, bipinnate to 3-pinnatifid; pinnae 1520 pairs, subopposite to alternate, stalk 619 mm, basal pinnae slightly reduced, narrowly triangular-ovate, 1416 Χ 3.84.2 cm, apex caudate, 1-pinnate to 2-pinnatifid; pinnules 810 pairs, anadromous, stalks 23 mm, basal pinnules about equal in size and shape, rhomboid, 1.52 Χ 11.5 cm, apex obtuse, base cuneate-truncate, almost symmetrical to asymmetrical, margin dentate. Veins obvious, raised adaxially, veins flabellately anadromous, not reaching margin. Costa green, with reduced scales or subglabrous, adaxially sulcate with broad supravascular ridge. Fronds papery, green, subglabrous; rachis dull grayish brown, with reduced scales or subglabrous, adaxially sulcate. Sori 38 per pinnule, basal on subtending vein, linear, 38 mm; indusia brownish, membranous, entire, opening toward costule or costa.
* Dense forests; ca. 1200 m. NW Yunnan (Dulongjiang Valley).
Asplenium yui is a distinctive species, similar to A. pseudolaserpitiifolium, known only from a restricted region. It is also close to A. nitidum Swartz but that taxon has more trullate and acute pinnules.
大盖铁角蕨 da gai tie jiao jue
Asplenium cavalerianum Christ; A. grandifrons Christ; A. latecuneatum Christ, nom. illeg. (based on same type as A. cavalerianum)[check (Bull. Gιogr. Bot. Mans 20: 173. 1909)]; A. viridissimum Hayata.
Plants 60100 cm tall. Rhizome erect, apex scaly; scales medium to dark brown, lanceolate, margin subentire. Fronds caespitose; stipe dull gray-brown to gray-green, semiterete, 2043 cm, adaxially sulcate, base scaly, becoming subglabrous near rachis; lamina triangular-ovate, 4570 Χ 1835 cm, apex acute, 2-pinnate to 4-pinnatifid; pinnae 1619 pairs, subopposite to alternate, stalked, basal pinnae slightly reduced, middle pinnae largest, narrowly triangular, 1119 Χ 48 cm, often slightly falcate, apex acuminate, base asymmetrical, acroscopic side truncate, basiscopic side cuneate, 2- or 3-pinnate; pinnules 813 pairs, anadromous, basal pinnules free and stalked, distal pinnae adnate to costa, basal acroscopic pinnule largest, ovate-triangular, 2.54 Χ 1.62.2 cm, base cuneate, asymmetrical and often decurrent along costule, apex obtuse to subacute; ultimate pinnules 25 pairs, basal acroscopic segment largest, ovate, 1215 Χ 68 mm, apex obtuse to truncate, base cuneate and decurrent on costule or free, margin dentate, teeth deltoid-triangular and often submucronate. Veins obscure, anadromous, rarely simple with distinct terminal hydathode on adaxial side, not reaching margin. Fronds herbaceous, green; rachis gray-green, semiterete, lamina with 24 cellular uniseriate hairs with apical gland or subglabrous, adaxially sulcate but with distinct supravascular ridge; costa pale brown to green, terete-semiterete, adaxially sulcate with distinct supravascular ridge, winged near its apex. Sori 125 per pinnule, median on subtending vein, subelliptic, ca. 4 mm; indusia yellowish brown, semi-elliptic, membranous, with thin hyaline margin with short 13-celled uniseriate hairs, rolling back at maturity, opening toward costule, persistent. Plant tetraploid: 2n = 144.
Streamsides in forests; 9002600 m. Fujian, Guizhou, Hunan, Sichuan, Taiwan, Xizang, Yunnan [Bhutan, N India, N Myanmar, Nepal, N Vietnam].
The lamina of Asplenium bullatum varies from 2-pinnate to 4-pinnatifid, with the more divided plant considered typical and recognized as A. bullatum var. bullatum (大盖铁角蕨(原变种) da gai tie jiao jue (yua bian zhong)). Bipinnate forms have been called A. bullatum var. shikokianum (Makino) Ching & S. H. Wu (FRPS 4(2): 75. 1999; 稀羽铁角蕨 xi yu tie jiao jue). However, A. Χshikokianum Makino (Bot. Mag. (Tokyo) 6: 45. 1892[TROP: Bot. Mag. (Tokyo) 13: 13. 1899??]; pro sp.) is the name for the natural hexaploid (Shimura & Matsumoto, [??citation]. 1976) hybrid between octoploid A. wrightii and A. ritoense Hayata (described from Japan), and cannot be used to describe the less divided forms of A. bullatum. Many specimens currently filed in Chinese herbaria as A. shikokianum or as A. bullatum var. shikokianum have good spores and are not hybrids. However, true A. Χshikokianum is not uncommon where its parents grow together (e.g., Guizhou, Hunan, Sichuan, and Taiwan). Asplenium Χwangii Kuo, the hybrid between A. bullatum and A. wrightii, is morphologically similar to A. Χshikokianum, and can easily be confused. Asplenium Χwangii Kuo was described from Taiwan (Kuo, [??citation]. 1998), and may be present where the parent taxa grow together (e.g., Fujian, Guizhou, Hunan, Sichuan, Taiwan, and Yunnan).
閩浙铁角蕨 min zhe tie jiao jue
Asplenidictyum wilfordii (Mettenius ex Kuhn) Nakai; Asplenium curtidens (Christ) Koidzumi; A. fengyangshanense Ching & C. F Zhang; A. wilfordii var. curtidens Christ; Tarachia wilfordii (Mettenius ex Kuhn) H. Itτ.
Plants 3040 cm tall. Rhizome erect, scaly; scales red-brown to black, narrowly triangular, 36 mm, entire. Fronds caespitose; stipe dull to semi-shiny, dark brown abaxially, green and sulcate adaxially, semiterete, 1225 cm, base with brown narrow scales; lamina ovate to narrowly triangular, 1030 Χ 513 cm, apex acute, 2-pinnate to 4-pinnate; pinnae 815 pairs, subopposite to alternate, with distinct stalk 38(10) mm, basal pair largest or slightly reduced, triangular-ovate, 38 Χ 24(6) cm, base asymmetrical, broadly cuneate, 2- or 3-pinnatifid, apex obtuse to acuminate-caudate; pinnules 46 pairs, anadromous, basal acroscopic pair largest, ultimate pinnules or segments 2 or 3 pairs, basal acroscopic pair largest, rhombic-cuneiform to linear, 412 Χ (1.5)45 mm, base cuneate and decurrent on costule, 2- or 3-partite, lateral side entire, apex serrate with linear to triangular teeth, apex obtuse to truncate. Veins distinct on both sides, raised adaxially, veinlets 14 per segment, not reaching margin. Fronds herbaceous to subleathery, dark green, paler abaxially, yellowish green to brown when dry; rachis dull to semi-shiny, dark brown abaxially becoming green toward apex, green adaxially with raised median ridge, semiterete, scaly; costa green, adaxially shallowly sulcate with prominently raised supravascular ridge, abaxially flat, with small fibrillar scales. Sori 13 per segment, linear to oval, 13 mm; indusia grayish white, membranous, entire, often opening toward each other. Plants tetraploid: 2n = 144.
On rocks in forests; 200300 m. Fujian, Jiangxi, Taiwan, Zhejiang [Japan, Korea].
The type is from Japan.
拟大羽铁角蕨 ni da yu ti jiao jue
Plants up to 1 m tall or more. Rhizome erect, apex scaly; scales dark brown, narrowly triangular, entire. Fronds caespitose; stipe dull to subshiny, purplish black to brown-gray, 2045 cm, adaxially sulcate, base with brown narrowly triangular scales, glabrous toward rachis; lamina triangular to ovate-elliptic, 4070 Χ 1530 cm, apex acute, 3-pinnate to 4-pinnatifid; pinnae up to 16 pairs, basal pinnae not reduced and subopposite, upper ones alternate, with stalk up to 1.5 cm, triangular to narrowly triangular, 715 Χ 411 cm, base broadly cuneate, bipinnate to tripinnatifid, apex acute; pinnules 1014 pairs, anadromous, stalks 57 mm, less than twice as long as wide, basal acroscopic and basiscopic pinnules about equal, basal pair largest, triangular to rhombic, 27 Χ 1.54 cm, base obliquely truncate, pinnate to 2-pinnatifid, apex acute; ultimate pinnules 26 pairs, anadromous, basal segment pair on basal pinnules subequal, largest, rhombic to cuneate, 1.52 Χ 1.51.7 cm, base broadly cuneate to cuneate, shortly stalked, 2- or 3-partite or simple, apex obtuse; segment rhombic to cuneate, lateral sides entire, apex bicrenate, apex obtuse to truncate. Veins distinctly raised adaxially, venation anadromous, veins almost reaching margin, pseudovein in first acroscopic sinus of basal pinnules. Fronds soft papery, brown-green when dry, subglabrous; rachis and costa dull purplish black to gray-brown, semiterete, adaxially sulcate; costule green. Sori linear, 46 mm; indusia membranous, entire, opening toward main vein of segment, persistent.
On rocks at streamsides in forests; 800900 m. Guangdong, Guangxi, Taiwan, Yunnan [Vietnam].
The type is from Guangxi (Shiwan Da Shan).
假大羽铁角蕨 jia da yu tie jiao jue
Asplenium neolaserpitiifolium Tardieu & Ching.
Plants up to l m tall. Rhizome shortly creeping to ascending, thick, apex scaly; scales pale to dark brown, narrowly triangular to linear, entire. Fronds subcaespitose to caespitose; stipe dull gray to gray-brown or almost black, 1540 cm, semiterete, adaxially sulcate, base scaly, upward subglabrous; lamina triangular-ovate, 1555(70) Χ (9)2540 cm, apex acute, tripinnate to 4-pinnatifid; pinnae 1015 pairs, subopposite to alternate, stalk up to 1 cm, basal pair largest or slightly reduced, triangular to narrowly triangular, often slightly falcate, 1025 Χ 615 cm, base broadly cuneate, 2-pinnate to 3-pinnatifid, apex acuminate to caudate; pinnules 1012 pairs, anadromous, stalk up to 4 mm, more than twice as long as wide, basal acroscopic and basiscopic pinnules about equal, narrowly triangular-ovate, 410 Χ 1.54.5 cm, base asymmetrical, acroscopic side truncate, basiscopically cuneate, pinnate to 2-pinnatifid at base of frond, apex acute to caudate; ultimate segments rhombic to obovate or oblong, 811 Χ 35.5 mm, base cuneate or segments adnate to costa, apex crenate to sinuate, with blunt teeth, apex obtuse. Costa dull gray to gray-brown abaxially, often becoming green toward apex, adaxially sulcate and often green, with gradually more prominent supravascular ridge toward apex; costule green, sulcate with median ridge adaxially; veins adaxially raised and obvious or flat and hardly visible, anadromous to subflabellately branching, not reaching margin; all axes with reduced scales to subglabrous. Fronds herbaceous to subleathery, dark green to grayish or brown-green after drying; rachis dull gray to brown-gray, semiterete, adaxially sulcate, with supravascular ridge and becoming green toward apex. Sori (1 or)24(7) per ultimate segment, linear, 36(8) mm; indusia pale yellow-brown, linear, membranous, entire, opening toward costule or main vein of segment, persistent. Spores pale brown with lophate (costate) perispore. Plants octoploid: 2n = 288.
On rocks at streamsides or epiphytic in forests; 1001100 m. Fujian, Guangdong, Guangxi, Hainan, Hunan, Taiwan, Xizang, Yunnan [India, Indonesia, Japan, Malaysia, Myanmar, Philippines, Thailand, Vietnam].
The type is from Vietnam.
Asplenium pseudolaserpitiifolium belongs to a group of large 3- to 4-pinnate species similar to A. laserpitiifolium Lamarck and revised by Tardieu and Ching ([??citation]. 1936), who described five new Indochinese species. Study of the types shows that characters used to describe or key out their taxa are variable and not diagnostic. Our field studies show that habitat conditions and plant age may strongly influence frond morphology and lamina texture. Plants collected after desiccation stress have frond segments ± curled inward and have a more narrowly cuneate form instead of the original rhombic shape. In the same paper, A. pseudolaserpitiifolium was described and keyed out as a terrestrial plant, but data on the labels of several paratypes state plants were epiphytic. Flow cytometric analyses show plants are octoploids in Guangdong, Guangxi, and Hainan; no data are available from the literature. Spores of the type specimens of A. neolaserpitiifolium and A. pseudolaserpitiifolium are identical.
Chinese pinyin
Plants 315 cm tall. Rhizome erect, scaly; scales dark brown, narrowly triangular, margin fimbriate at base. Fronds caespitose; stipe purplish black to castaneous, semi-shiny, 0.54 cm, adaxially sulcate, with dark brown narrowly triangular scales mixed with filiform scales; lamina linear or linear-lanceolate, 315 Χ 0.52.5 cm, attenuate to both ends, apex pinnatipartite or long flagelliform with terminal budlet, 1-pinnate to 2-pinnatifid; pinnae 1020 pairs, pinnae sessile or shortly stalked, basal pinnae usually smaller and flabellate, middle pinnae largest, distinctly separated from each other, linear-elliptic, (2)510(15) Χ 1.55(7) mm, base almost symmetrical to asymmetrical, acroscopic side truncate, basiscopic side broadly cuneate, pinnatipartite, apex obtuse and with one gemma in apical sinus; segments 14 pairs, apex obtuse but with 2 or 3 teeth, basal acroscopic segment not much enlarged. Veins not prominent, anadromously forking, not reaching margin. Fronds herbaceous, green, lamina with uniseriate gland-tipped hairs or subglabrous, average guard cell length 3439 ΅m; rachis with purplish brown to castaneous color extending from basal part up to 1/22/3 its length on abaxial side, apical part green or stramineous when dry, with purplish black hairlike scales, adaxially green or stramineous, flat or slightly sulcate, green. Sori usually one per segment, often confluent at maturity, subelliptic, 0.31 mm, basal to submedial on subtending veinlets, close to costa; indusia grayish green to grayish brown, subelliptic, membranous, repand to entire, mainly opening toward costa. Spores with lophate (costate-cristate) perispore, average exospore length 2428 ΅m. Plants sexual, diploid: 2n = 72.
In rock (limestone) crevices in forests; 11002800 m. Sichuan, Yunnan [Nepal, Vietnam].
Asplenium lushanense is the diploid ancestor of the autotetraploid A. exiguum (Viane & Reichstein, [??citation]. 2003; Wang, [??citation]. 2003), see below.
云南铁角蕨 yun nan tie jiao jue
Asplenium fontanum (Linnaeus) Bernhardi var. exiguum Beddome; A. fontanum var. yunnanense (Franchet) Beddome; A. kangdingense Ching & H. S. Kung; A. loherianum Christ; A. moupinense Franchet; A. moupinense var. dareiforme Franchet [dareaeformis]; A. woodsioides Christ; A. yunnanense Franchet; A. yunnanense var. dareiforme (Franchet) H. S. Kung [daraeforme].
Plants 525 cm tall. Rhizome erect, scaly; scales dark brown to black, narrowly triangular, margin fimbriate at base. Fronds caespitose; stipe 15 cm, abaxially purplish black to reddish brown and semi-shiny, adaxially sulcate, with blackish brown narrowly triangular scales mixed with filiform scales; lamina lanceolate, 315 Χ 0.82.5(4) cm, attenuate to both ends, apex pinnatipartite or shortly flagelliform and rooting, 1- or 2-pinnate, up to 3-pinnatifid in large specimens; pinnae 1020 pairs, shortly stalked, basal pinnae opposite and often strongly reduced or flabellate, upward becoming alternate; middle pinnae largest, narrowly triangular to ovate-elliptic or oblong, (2)1015(20) Χ 27 mm, base almost symmetrical to asymmetrical, acroscopic side truncate, basiscopic side cuneate, pinnatisect to pinnatifid, apex obtuse and with one gemma in apical sinus; segments 24 pairs, apex obtuse and dentate-serrate with 24 teeth; teeth obtuse, mucronate or acute. Veins not prominent, anadromously forking, not reaching margin. Fronds firmly herbaceous, green, lamina with uniseriate gland-tipped hairs or subglabrous, average guard cell length 4350 ΅m; rachis with purplish brown to castaneous color extending from basal part up to 2/3 its length on abaxial side, apical part green or stramineous, with purplish black hairlike scales, adaxially green or stramineous, flat or slightly sulcate, green. Sori usually 24 on basal acroscopic segment, on others one per segment, often confluent at maturity, basal to submedial on subtending veinlets, close to costa, subelliptic to elongate, ca. 1 mm; indusia grayish green to grayish brown, narrowly elliptic, membranous, repand to entire, opening toward costa or major vein. Spores with lophate (costate-cristate) perispore, average exospore length 3135 ΅m. Plants sexual, autotetraploid: 2n = 144.
In crevices of (limestone) rocks in forests; 11003300 m. Guangxi, Guizhou, Hebei, Henan, Hunan, Shanxi, Sichuan, Taiwan, Xizang, Yunnan [N India, Mongolia, N Myanmar, Nepal, Philippines, Russia (SW Siberia), Thailand, N Vietnam; North America (Mexico, USA)].
The type is from S India.
Large specimens of Asplenium exiguum are similar to well-developed plants of A. nesii, and can best be distinguished by the presence of gemmae in the apical notch of the pinnae.
Asplenium exiguum is an autotetraploid taxon that most probably originated via chromosome doubling in diploid A. lushanense. It is a variable species with an easily overlooked, tiny bud in the sinus at the pinna apex, and well-illustrated in Ching ([??journal or book citation]: pl. 174: 2b. 1937). The morphological variability, both within China and throughout the Himalaya, is relatively large, though mainly restricted to size differences. Growing conditions may strongly influence frond length and the shape of pinnae (from almost oblong to triangular). In some populations (S India, China, Philippines), the rachis is flagelliform and terminates in a slender tail with a terminal budlet. This variation led to much confusion and the description of several (local) species, e.g., Asplenium glenniei Baker, A. loherianum, A. moupinense, A. woodsioides, and A. yunnanense. Asplenium moupinense var. dareiforme Franchet (1887) represents a luxurious, well developed plant that does not differ in any essential character from other members of this group. Since there are no fundamental morphological, nor cytological, differences between these taxa, we treat them as synonyms of tetraploid A. exiguum following Hope ([??citation]. 1899; [??citation]. 1901), Ching ([??citation]. 1937), Tagawa ([??citation]. 1939), Tagawa and Iwatsuki ([??citation]. 1985), and Mickel and Smith ([??citation]. 2004). The diploid and tetraploid species within this complex can be separated by microcharacters (Viane & Reichstein, [??citation]. 2003): Asplenium lushanense (diploid) with an average exospore length of 2428 ΅m and stomates 3240 ΅m, and tetraploid A. exiguum with average exospore length of 3036 ΅m and stomates 4352 ΅m. Where both species grow together, their sterile triploid hybrid, with aborted spores, an intermediate morphology, and often showing hybrid vigor, is common. Asplenium Χmickelii Viane, Reichstein, hybr. nov. Type: Mexico. Oaxaca: Destrito Ixtlαn. 2 km W of Ixtlαn de Juαrez on Rd 175, 1900 m, 16 Sep 1972, J.T. Mickel & L. Pardue 6488-A (left hand specimen) (holotype, NY; merotype: TR-4095, raised from frond gemmae of the holotype, GENT). Planta hybrida, inter parentes A. exiguum et A. lushanensem intermedia, sporis abortivis, numero chromosomatico triploideo 2n = 108, meiosi 04 trivalentibus, 3236 bivalentibus 2936 univalentibusque differt.
虎尾铁角蕨 hu wei tie jiao jue
Asplenium elegantulum Hooker.
Plants 1030 cm tall. Rhizome short, erect to shortly creeping, occasionally gemmiferous, apex scaly; scales dark brown to black, narrowly triangular, margin fimbriate with short glandular hairs or entire. Fronds caespitose; stipe 410 cm, semiterete, basal part abaxially shiny castaneous but upward becoming green, adaxially green and sulcate, with dark brown hairlike scales or subglabrous; lamina lanceolate, 1027 Χ 24(5.5) cm, reduced at both ends, apex acute to acuminate, 1- or 2-pinnate; pinnae 1222 pairs, subopposite to alternate, shortly stalked, lower pinnae gradually reduced, deltoid to semi-orbicular, less than 5 mm, middle pinnae narrowly triangular, 12 Χ 0.61.2 cm, apex acute, pinnatipartite to 1-pinnate, its upper segments gradually more adnate to costa; pinnules or segments 46 pairs, anadromous, alternate, basal acroscopic pair largest, elliptic or ovate, 47 Χ 35 mm, base cuneate, free or decurrent on costa, margin serrate, apex obtuse. Costa distinct, veins obscure, anadromously 2-forked or simple, terminal hydathode distinct and not reaching margin. Fronds herbaceous, often dimorphic and sterile fronds small, green, subglabrous; rachis green or stramineous when dry or base shiny castaneous abaxially, subglabrous, adaxially sulcate, apical part with narrow lateral wings. Sori median to submedian on veinlet, linear, ca. 1 mm; indusia grayish, semi-elliptic, membranous, entire to repand, opening toward costa. Plants sexual, diploid: 2n = 72.
On wet rocks in forests or on man-made constructions[??buildings or walls]; 1001600 m. Anhui, Fujian, Gansu, Guangdong, Guizhou, Hebei, Heilongjiang, Henan, Hunan, Jiangsu, Jiangxi, Liaoning, Shaanxi, Shandong, Shanxi, Sichuan, Taiwan, Yunnan, Zhejiang [Japan, Korea, Russia].
The type is from Japan.
Asplenium incisum is an ancestral diploid (Kurita, [??citation]. 1962; [??citation]. 1966) that hybridized with A. ruprechtii to form a diploid sterile hybrid, which via polyploidy led to the tetraploid A. castaneo-viride (= A. kobayashii).
西北铁角蕨 xi bei tie jiao jue
Asplenium barkamense Ching; A. tianshanense Ching.
Plants (2)612(25) cm tall. Rhizome erect, short, apex scaly; scales dark brown to black, narrowly triangular, 2.53 mm, entire. Fronds caespitose; stipe 2.58(10) cm, basal part semi-shiny, castaneous or dark brown to black, toward rachis green or stramineous, sparsely covered with small, black, hairlike scales, adaxially sulcate but compressed when dry; lamina lanceolate, 46(16) Χ 12(5) cm, reduced at base, 2- or 3-pinnatifid; pinnae 79(15) pairs, alternate or subopposite, shortly stalked, middle pinnae largest, elliptic, 515(25) Χ 38(15) mm, base subsymmetrical to asymmetrical, truncate to cuneate, 1- or 2-pinnatifid in large plants, apex acute and pinnatifid, without gemma; pinnules or segments 35(7) pairs, alternate, basal acroscopic pinnule largest, oblong to linear, 46 Χ 12(4) mm, apex obtuse, base cuneate, decurrent and adnate to costa, margin serrate-dentate, teeth often long, obtuse to subacute; other pinnules or segments smaller, many adnate to costa. Veins obscure, 2-forked or simple, not reaching margin. Fronds herbaceous to subleathery, gray-green; rachis green or stramineous when dry, rarely brown at base abaxially, adaxially sulcate, with small, dark brown, hairlike scales. Sori 24 per pinnule, basal to submedian on subtending vein and thus close to costa or costule, confluent at maturity, linear, 11.5 mm; indusia grayish brown to whitish, linear-elliptic, membranous, entire, opening toward costa or costules. Spores with lophate (costate) perispore. Plants tetraploid: 2n = 144.
Crevices of relatively dry rocks; 10004000 m. Gansu, Hebei, Nei Mongol, Ningxia, Qinghai, Shaanxi, Shanxi, Sichuan, Xinjiang, Xizang, Yunnan [Afghanistan, N India, N Pakistan].
The type is from Shaanxi (Shanyang).
Asplenium nesii is similar to larger forms of A. exiguum, but their pinnae apices are without gemmae, the stipe is green toward the rachis, and the rachis itself is usually entirely green. Asplenium barkamense is a luxurious form from Sichuan, but locally not rare in Xizang. Type specimens of A. Χmainlingense Ching & S. K. Wu (Fl. Xizang 1: 184. 1983; pro sp.) are hybrids with aborted spores. They probably represent a cross between A. nesii and A. varians or A. tenuicaule, which all occur in the area close to the locus typicus near Mainling, Xizang.
西藏铁角蕨 xi zang tie jiao jue
Asplenium pseudofontanum Kossinsky, Not. Syst. Herb. Hort. Petrop. 3: 122. 1922.
Plants (5)1320 cm tall. Rhizome erect, short, apex scaly; scales dark gray-brown, narrowly ovate-triangular, 34 Χ 0.50.6 mm. Fronds caespitose; stipe (1)48 cm, base semi-shiny, castaneous to dark brown for 0.22 cm on adaxial side, abaxially often whole stipe brown, green toward apex, base scaly, subglabrous toward rachis; lamina lanceolate, 810(15) Χ 13 cm, widest slightly above middle, gradually tapering to both ends, bipinnate to tripinnatifid at base, apex acute; pinnae 1020(30) pairs, subopposite to alternate, stalk 0.21 mm, lower pinnae reduced, sessile and more remote, middle pinnae narrowly ovate to triangular, 512(25) Χ (2)58(12) mm, pinnatisect to pinnate, apex acute, base broadly cuneate, subsymmetrical, basal acroscopic pinnule largest, broadly ovate, 24(7) mm, close to or covering rachis, anadromous; pinnules 35(7) pairs, with triangular-ovate lobes, 45 Χ 34 mm, base broadly cuneate and decurrent on short stalk, palmate or subpinnatilobate, apex obtuse; segments elliptic, ca. 1 mm, teeth mucronate or acute. Veins obvious, raised adaxially, one per segment, not reaching margin. Fronds herbaceous, pale green to grayish green when dry, lamina subglabrous, average stomatal guard-cell length 3038 ΅m; rachis semi-terete, green, or occasionally brown at base abaxially, adaxially sulcate. Sori 13 per pinnule, median on acroscopic veins, linear to oval, 11.5 mm; indusia whitish green or hyaline, semi-elliptic, membranous, entire, opening toward costa or costule. Spores lophate (reticulate), average exospore length 3035 ΅m. Plants sexual, diploid: 2n = 72.
On limestone rocks in forests; 15002100 m. Xinjiang [N Afghanistan, N India, E Kazakhstan, E Kyrgyzstan, Nepal, N Pakistan, E Tajikistan, Turkmenistan, Uzbekistan; SW Asia (Iran)].
The type is from Tajikistan.
Asplenium fontanum consists of two morphologically slightly different but geographically widely separated, diploid subspecies with homologous chromosomes (Reichstein & Schneller, [??citation]. 1982; Gibby, [??citation]. 1982). The typical subspecies, A. fontanum subsp. fontanum, only grows on limestone rocks in southern and central Europe. In Asia, it is replaced by this slightly different subspecies with somewhat wider fronds, deeply incised pinnae, and narrower segments. Its perispore is also slightly different with an outer perisporal layer with numerous small holes (fenestrate-reticulate).
线柄铁角蕨 xian bing tie jiao jue
Asplenium varians Wallich ex Hooker & Greville var. sakuraii Rosenstock.
Plants 38(13) cm tall. Rhizome erect, short, apex scaly; scales dark brown to black, triangular-ovate, margin shortly fimbriate glandular. Fronds caespitose; stipe green, slender, threadlike, 0.32.5(9) cm, subglabrous, adaxially sulcate; lamina triangular to narrowly triangular, (0.6)1.56 Χ 0.52.5 cm, apex obtuse to subacute, 2-pinnate to almost 3-pinnate; pinnae (3)57 pairs, alternate or basal ones subopposite, stalk slender, threadlike, often at least one stalk with tiny gemma in acroscopic axil, basal pinnae not reduced, broadly ovate to triangular, 4(12) Χ 4(6) mm, pinnate to 2-pinnatifid, upper pinnae trifid to bifid; pinnules 2 or 3(or 4) pairs or 1 pair and a trifid apex, usually bifid, trifid, or simple, basal acroscopic pinnule largest, cordiform or elliptic, 23 Χ 12 mm, base cuneate, decurrent on costa, margin entire, apex variable, obtuse or mucronate to acute; pinnae near apex simple or bifid. Veins obscure, anadromously pinnate, simple, one per segment or pinnule, with hydathode at end, not reaching margin. Frond herbaceous, green; rachis green, subglabrous, adaxially shallowly sulcate, apex not flagelliform, lamina subglabrous, average guard cell length 4549 ΅m. Sori one per segment or pinnule, basal to median on subtending vein, subelliptic, 12 mm; indusia grayish green, elliptic, membranous, entire, opening toward costa or costule, persistent. Sporangia with 32 spores, perispore lophate (alate), average exospore length 3842 ΅m. Plants sexual, diploid: 2n = 72.
Moss cushions on shaded, wet limestone rocks and crevices; 18003000 m. ?Gansu, Guizhou, Hunan, ?Shaanxi, Sichuan, Taiwan, Yunnan [Bhutan, India, Japan, Korea, Nepal].
The type is from Japan.
Asplenium capillipes is the smallest species of the genus in China, and may be overlooked since it often grows hidden among mosses. It is reported from Gansu (Tianshui), Hunan, and Shaanxi (Tibia Shan), but we have seen no specimens; however, it should be noted that large plants of this species are similar to small plants of A. tenuicaule and may be confused with it. Identification of vouchers from Bhutan (Itτ, [??citation]. 1966) needs verification since some belong to A. fugax, which has a flagelliform apex in contrast to A. capillipes. Plants growing in thick moss cushions may have stipes up to 9 cm. Asplenium capillipes is diploid with n = 36II during meiosis (Mitui, [??citation]. 1970) and 2n = 72 chromosomes in vegetative cells. It is remarkable that it has only 32 relatively large spores per sporangium.
Chinese pinyin
Type: China. Yunnan: shaded rocks in forest of San-tcha-ho, above Mo-so-yn, 17 May 1887, Delavay s.n. sheet 1, plant 7 (lectotype, designated here, P!).
Plants 38 cm tall. Rhizome erect, short, scaly; scales dark brown to black, narrowly triangular, margins fimbriate. Fronds caespitose; stipe green, thin, threadlike, 0.51.5(2.5) cm, subglabrous, adaxially shallowly sulcate; lamina narrowly triangular to linear, 26 Χ 0.71.2 cm, base slightly or not reduced, bipinnate, apex acute or truncate, usually flagelliform with terminal small bud; pinnae 613 pairs, alternate or subopposite, shortly stalked and only occasionally with a gemma in upper axil, ovate, 46 Χ 34 mm, pinnate; pinnules 1 or 2 pairs, usually simple, elliptic, 23 Χ 12 mm, base cuneate, margin entire, apex acute; pinnae near apex less divided. Veins simple, one per pinnule or segment, not reaching margin. Fronds thinly herbaceous, green to grayish green when dry; rachis green, subglabrous, adaxially sulcate. Sori one per segment or pinnule, subelliptic, 0.51 mm; indusia grayish green, subelliptic, membranous, entire, opening toward costa, persistent. Sporangia with 32 spores, perispore lophate (alate), average exospore length 4448 ΅m.
In deep crevices of shaded (limestone) rocks; 25003500 m. Guizhou, Sichuan, Yunnan [Bhutan, Nepal].
The type is from China.
Asplenium fugax is easily overlooked since it is a small species hidden in moss cushions under boulders. Plants are often identified or reported as A. capillipes (e.g., Itτ, [??citation]. 1966; Nakaike, [??citation]. 1986), but that taxon never has a flagelliform apex. Christ ([??citation]. 1905) cited two syntypes: one collection by Delavay and another by Farges, both kept in Paris (P). Delavays gathering is a mixture mounted on three separate sheets numbered 1, 2, and 3. Most plants fit Christs description and belong to A. fugax, but there is also a Woodsia specimen on sheet 2. Plant 7 on sheet 1 (bearing Delavays original label) is well pressed, the most fertile and best developed, and it is selected here as lectotype. The other syntype [Sichuan, Tchen-kιou-tin, 2000 m, August, Farges 657 (P!)] belongs to A. capillipes.
细裂铁角蕨 xi lie tie jiao jue
Asplenium concinnum Wallich, nom. nud.; A. tenuissimum Hayata.
Plants (10)2045 cm tall. Rhizome erect, short, apex scaly; scales dark brown, triangular, margin dentate-fimbriate or entire. Fronds caespitose; stipe semiterete, 721 cm, abaxially green or castaneous up to middle, with scales or subglabrous, adaxially shallowly sulcate with supravascular ridge; lamina triangular to ovate, 1230 Χ 613 cm, apex acute, 2- or 3-pinnate or up to 4-pinnatifid; pinnae 1016 pairs, alternate, stalk short, basal pair almost unreduced, narrowly triangular, 38 Χ 1.63 cm, base truncate to cuneate, up to tripinnatifid, apex acute-acuminate; pinnules (4)612(16) pairs, alternate, anadromous, shortly stalked, usually acroscopic segment larger, ovate, 1.32.8 Χ 11.3 cm, up to 2-pinnatifid; secondary pinnules 3 or 4 pairs, 58 Χ 46 mm, base cuneate and decurrent on costule, 24-partite, apex obtuse; ultimate segments ovate-lanceolate, ca. 1.5 mm wide, with a single vein, apex acute. Costa slender, adaxially sulcate with supravascular ridge to almost flat when dry, terminal hydathode obvious, not reaching margin. Fronds thinly herbaceous, green; rachis green, adaxially shallowly sulcate with median supravascular ridge or flat, adaxial side of costae or costules with small gemma or plantlet. Sori one per ultimate segment, median on subtending vein, elliptic, 1.52 mm; indusia whitish to grayish green, linear to elliptic, membranous, entire, opening toward margin costule or central veinlet, persistent. Spores with lophate (alate) perispore. Plants sexual, diploid or tetraploid: 2n = 144.
On wet rocks in mixed forests; 8002900 m. Guangxi, Guizhou, Hainan, Hunan, Sichuan, Taiwan, S Xizang, Yunnan [Bhutan, India, Indonesia, Malaysia, Myanmar, Nepal, Philippines, Sri Lanka, Thailand, Vietnam].
The type is from Nepal.
Asplenium tenuifolium is common in the Himalaya. It is probably an aggregate of diploid plants with 2n = 72 chromosomes (reported from India; Bir, [??citation]. 1960; [??citation]. 1965) and tetraploids (specimens from Guangxi and Guangdong). More studies are needed. Most plants have small dormant buds (one or more) on the adaxial side of the costa (near the first acroscopic costule or more distal) or on the costules. It is often fertile when still young (var. minor S.H. Wu) and can then be confused with A. capillipes (gemmae in axil between costa and rachis) or with A. tenuicaule (without gemmae). The presence and position of its gemmae are diagnostic. We agree with Tagawa (1939) that there are no significant differences between A. tenuissimum and A. tenuifolium.
圆叶铁角蕨 yuan ye tie jiao jue
Asplenium ruta-muraria Linnaeus subsp. dolomiticum Lovis & Reichstein, Brit. Fern Gaz. 9: 143. 1964; A. deqenense Ching; A. suborbiculare Ching.
[??add info to make description parallel to others] Morphologically identical to Asplenium ruta-muraria, but with average exospore length 3540 ΅m. Plants sexual, diploid: 2n = 72.
In rock crevices (often dolomite); 15003300 m. Hebei, Sichuan, Xizang, Yunnan [Afghanistan, Mongolia, Russia (E Siberia); SW Asia (Iran, Turkey), S Europe].
Contrary to previous reports, new and more extensive research in southern Europe has shown that there are no gross morphological differences between Asplenium dolomiticum and A. ruta-muraria (see above). Asplenium deqenense and A. suborbiculare, both with an average exospore length of 3940 ΅m, are considered juvenile, depauperate forms of A. dolomiticum. Small juvenile forms with simple or ternate fronds are not rare in populations of both A. ruta-muraria and A. dolomiticum.
卵叶铁角蕨 luan ye tie jiao jue
Asplenium ruta-muraria var. subtenuifolium Christ; A. subtenuifolium (Christ) Ching & S. H. Wu; Tarachia ruta-muraria (Linnaeus) C. Presl.
Plants 210(15) cm tall. Rhizome erect to shortly creeping, scaly; scales blackish brown, narrowly triangular, occasionally with marginal glands, glands also on apex, subentire. Fronds caespitose; stipe (0.5)25(6) cm, green or grayish green to stramineous when dry, base castaneous to dark purplish or blackish brown, with numerous short (4-cellular) glandular hairs and hairlike scales, subglabrous toward rachis; lamina deltoid-triangular to ovate, (1)25(8) Χ 13(5) cm, apex obtuse to subacute, 2-pinnate or pinnate-pinnatifid; pinnae 14 pairs, subopposite or alternate, basal pinnae largest, (4)812 Χ (4)810 mm, stalked, pinnae triangular, apex obtuse, imparipinnate or ternate with up to 3 lateral segments, segments often similarly divided; ultimate segments broadly flabellate to rhombic-ovate or trapeziform, (3)46 Χ 25 mm, base broadly cuneate and decurrent on costa, margin irregularly dentate, apex obtuse. Veins obscure, flabellately anadromous, and almost parallel. Fronds subleathery, grayish green when dry; rachis and costa green, with (partly) deciduous glandular hairs, subglabrous when old, adaxially sulcate, when dry often flat. Sori 512 per pinna, median on subtending vein, confluent at maturity, subelliptic to linear, 15 mm; indusia gray-glaucous to brown, linear, thinly membranous, margin with long hairs, opening toward major veins or costa, concealed by sporangia at maturity. Perispore lophate (costate-cristate), average exospore length 4149 ΅m. Plants sexual, autotetraploid: 2n = 144.
On limestone rocks; 8002700 m. Gansu, Guizhou, Hunan, Liaoning, Nei Mongol, Shaanxi, Shanxi, Sichuan, Xinjiang, Yunnan [Afghanistan, Himalaya[please list countries], India, Japan, Kazakhstan, Korea, Kyrgyzstan, Nepal, Pakistan, Russia, Tajikistan; NW Africa, Europe, North America].
The type is from Europe.
Asplenium ruta-muraria is an autotetraploid species, very common and widespread in Europe, where it often grows on man-made constructions[??buildings, walls]. It has originated by chromosome doubling in its diploid ancestor A. dolomiticum (see below), from which is genetically isolated, and which has smaller spores and half the number of chromosomes. Their triploid sterile hybrid, A. Χbaldense Sleep et al., is usually found where both species grow together. There is considerable variation in frond shape, and size and degree of dissection, best known and mainly described from Europe (e.g., von Heufler, [??citation]. 1856; Luerssen, [??citation]. 1889; Christ, [??citation]. 1900; [??citation]. 1903; Rosenstock, [??citation]. 1902; Fiori, [??citation]. 1943), and illustrated in Christ ([??citation]. 1903), Ogata ([??citation]. 1930), Khullar ([??citation]. 1994), and Wu ([??citation]. 1999). A hexaploid is known from Japan (Reichstein, [??citation]. 1981).
贵阳铁角蕨 gui yang tie jiao jue
Asplenium cuneifolium Viviani var. vegetius Christ; A. interjectum var. elatum Christ; A. longkaense Rosenstock.
Plants (10)2030 cm tall. Rhizome erect, short, apex scaly; scales black, linear, entire. Fronds caespitose; stipe 1218 cm, semiterete, abaxially shiny castaneous, often green toward rachis, subglabrous, adaxially sulcate and with median ridge; lamina broadly triangular to deltoid, 817 Χ 57 cm, apex acuminate-caudate, bipinnate to tripinnate at base; pinnae 57 pairs, alternate or basal pinnae subopposite, long stalked, basal pair largest, triangular, 49 Χ 36 cm, base truncate, subsymmetrical, 1- or 2-pinnate, apex acute to acuminate-subcaudate; pinnules (1 or)2 or 3(or 4) pairs, alternate, anadromous, stalked; basal pinnules largest, ovate-triangular, 1.53 Χ 12.6 cm, base cuneate, pinnatisect, apex obtuse to acute; ultimate pinnules 1 or 2 pairs, basal acroscopic segment largest, ovate-oblong, 614 Χ 49 mm, base cuneate and decurrent on costule, margins serrate, apex obtuse. Costa and basal part of veins raised adaxially, flat abaxially, veins anadromous-subflabellately branching, terminal hydathode visible, not reaching margin. Fronds herbaceous, green; rachis semiterete, abaxially green or shiny castaneous up to middle, subglabrous, adaxially sulcate with green lateral wings and median supravascular ridge. Sori 24 per ultimate segment, median on veinlets, linear, up to 6 mm; indusia gray to brownish, linear, membranous, repand to entire, opening toward costa or costule.
In rock crevices in forests; 700900 m. Guizhou, Yunnan [N Thailand, N Vietnam].
The type is from Guizhou (Guiyuang, Qingyan).
细茎铁角蕨 xi jing tie jiao jue
Plants (2.5)615(25) cm tall. Rhizome erect, short, apex scaly; scales brown to dark brown, triangular to narrowly triangular, 13.5 Χ 0.30.7 mm, base fimbriate or entire. Fronds caespitose; stipe (0.7)1.55(10) cm, green but base castaneous, brown color occasionally extending into rachis, with small dark brown, fimbriate scales and uniseriate hairs, subglabrous when old, adaxially sulcate; lamina triangular to narrowly triangular-ovate, (2)49(16) Χ (0.8)1.23(6.5) cm, base broadly cuneate and slightly reduced, rarely pinnate-pinnatifid, usually bipinnate but occasionally up to 4-pinnatifid, apex acute-acuminate; pinnae (5)710(15) pairs, alternate or opposite, stalk narrow and up to 3 mm, lower pinnae usually slightly reduced, largest pinnae triangular-ovate, (4)614(26) Χ (3)59(18) mm, rarely up to 45 Χ 25 mm, base asymmetrical, acroscopic side truncate, basiscopic side cuneate, 1- or 2- pinnate, rarely up to 3-pinnatifid, apex obtuse; pinnules 24 pairs (rarely up to 7 pairs), alternate, anadromous, free or adnate, basal acroscopic segment largest and often pinnatifid to almost 2-pinnate in basal part of large fronds, fan-shaped or ovate to triangular-ovate, margin serrate to crenate with acute to mucronate teeth, apex obtuse. Veins obscure, veinlets 1- or 2-forked, not reaching margin. Fronds thinly herbaceous, green to gray-green or brown when dried, rachis green or occasionally with castaneous base (abaxially), adaxially sulcate, subglabrous, lamina with 3- or 4-celled uniseriate gland-tipped hairs, average guard cell length 4145 ΅m. Sori 13(or 4) per pinnule, median on acroscopic veinlets, occasionally confluent at maturity, oval-linear, (0.8)1.12(3) mm; indusia whitish to pale brown, oval-linear, membranous, entire to repand, opening toward major veins or toward costa. Spores with lophate or reticulate perispore, average exospore length 2832 ΅m. Plants sexual diploid: 2n = 72.
On trees or rocks in forests; 2003600 m. Chongqing, Gansu, Guizhou, Hebei, Heilongjiang, Henan, Hunan, Jiangsu, Jiangxi, Jilin, Liaoning, Nei Mongol, Qinghai, Shaanxi, Shandong, Shanxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Bhutan, India, Japan, Korea, Nepal, Pakistan, Philippines, Russia (S Siberia), Thailand; E Africa, Polynesia (Hawaii)].
Asplenium tenuicaule is an ancestral diploid species involved in a reticulate pattern of relationships unraveled in the 19801990s (Sleep & Reichstein, [??citation]. 1984; Viane & Reichstein, [??citation]. 2003) by micromorphological studies and cytological research in China (Y.-X. Lin, Z.-R. Wang), England (A. Sleep), Germany (H. Rasbach), and Switzerland (T. Reichstein, R. Viane). This reticulate evolution, with its inherent general morphological fuzziness, has led to much misunderstanding and confusion. Asplenium tenuicaule is parental to the following taxa with which it can easily be confused: A. kansuense, A. varians, A. altajense, A. anogrammoides, and A. kukkonenii. During sporogenesis, this species produces a relatively high number of diplospores, which may have played a role in the production of polyploid offspring, e.g., A. kansuense, A. mae (see below).
This is a widespread and variable species ranging from the eastern African highlands (rare) via the Himalaya to southern (Altai) and southeastern Siberia, Korea, China, Japan, and Hawaii. It can be divided into three varieties, with overlapping ranges in China and Japan. Few but fertile intermediates have been found. Hybridization experiments have shown that the varieties are genetically very similar (Viane & Reichstein, [??citation]. 2003). Kuo ([??citation] 1985) was probably the first to apply the name correctly.
1a. Perispore lophate and
middle perisporal layer reduced or absent between folds
.................................................................................................................... 70b.
var. subvarians
1b. Perispore reticulate or if lophate then middle perisporal layer present between folds of outer perispore
2a. Pinnae with marginal teeth rounded to mucronate, outer perispore a perforated reticulum without distinct ridges .......................................................................................... 70a. var. tenuicaule
2b. Pinnae with marginal teeth acute, outer perispore lophate with rounded ridges (costate-cristate) 70c. var. argutum
细茎铁角蕨(原变种) xi jing tie jiao jue (yuan bian zhong)
Asplenium borealichinense Ching & S. H. Wu; A. hebeiense Ching & S. H. Wu.
Marginal teeth of pinnae obtuse to (sub)mucronate. Spores with a typical reticulate perispore, consisting of a thin basal endoperispore surrounding smooth exospore, an extensive middle perispore of rodlets (collumellae) subtending outer perisporal network of wide pori (13 ΅m) but only faint or no crests.
On trees or rocks in forests; 2002900m. Hebei, Shaanxi, Shanxi, Sichuan, Taiwan, Xizang, Yunnan [Japan, Korea, Thailand; E Africa, Polynesia (Hawaii)].
The type is from China (Taiwan).
See illustrations in Y. L. Zhang et al. (Sporae Pterid. Sin. t. 77 bc, t. 81 cd; Pl. 51: 1920; Pl. 52: 1718[20?]. 1976).
The following names belong here but were not validly published because no Latin description or diagnosis was provided (Vienna Code, Art. 36.1): Asplenium hopehense Ching ([in Y. L. Zhang et al., Sporae Pterid. Sin. 249. 1976??]) and A. miyunense Ching and A. pseudovarians Ching (Fl. Beijing 1: 30 1984; 1995).
钝齿铁角蕨 dun chi tie jiao jue
Asplenium subvarians Ching in C. Christensen, Index. Filic. 3: 38. 1934, not A. fauriei Christ (1899); Anogramma fauriei Christ; Asplenium chengkouense Ching ex H. S. Kung; A. conmixtum Ching, [conmixum;]; A. siobarense Koidzumi, nom. superfl. (1935); A. tianmushanense Ching; Gymnogramma fauriei Christ.
Marginal teeth of pinnae mucronate to acute. Spores with lophate (cristate) perispore, middle perispore lacking in areolae between outer perispore folds, with few, narrow (less than 1 ΅m) pori [see illustration of pattern in e.g., Y. L. Zhang et al., Spore Morphology of Chinese Pteridophytes (1976, 1990): t. 77 jl; Pl. 51: 3, 21, 33)][Different plate numbers in 1976 original. Change to Sporae Pterid. Sin. Pl. 51: 2223; Pl. 52: 2728. 1976; Spore Morphol. Chin. Pteridophytes, t. 77 jl; Pl. 51. 3, 21, 33, 1990??].
On rocks in forests; 6003100 m. Chongqing, Gansu, Guizhou, Hebei, Henan, Heilongjiang, Henan, Hunan, Jiangsu, Jiangxi, Jilin, Liaoning, Nei Mongol, Qinghai, Shaanxi, Shandong, Shanxi, Sichuan, Xizang, Yunnan, Zhejiang [Bhutan, India, Japan, Korea, Nepal, Pakistan, Philippines, Russia (S Siberia)].
The type is from Japan.
Asplenium tenuicaule var. subvarians has often been mentioned under A. anogrammoides auct. non Christ (e.g., in Komarov, [??citation]. 1916; Fomin, [??citation]. 1930; [??citation]. 1934), a mistake caused by Christensens interpretation of A. anogrammoides based on wrong type material. Small specimens of this species can be similar to A. capillipes Makino, but that species usually has a small bud on the stalk of one of its basal pinnae and also has larger spores with an alate perispore. A narrow form of A. tenuicaule var. subvarians grows in the Altai mountains and is often misidentified either as A. anogrammoides auct. non Christ, or as A. altajense auct. non Grubov. A luxurious form was described as A. chengkouense. In the western Himalaya, plants tend to have less mucronate, but more slender and sharper teeth. Due to confusion with similar taxa, the distribution of this taxon is not well known.
尖齿铁角蕨 jian chi tie jiao jue
Replaced synonym: Asplenium argutum Ching, Acta Phytotax. Sin. 23(1): 9. 1985, not Asplenium argutum Kaulfuss, Enum. Filic. 176. 1824.
Pinnae with acute marginal teeth, Spores with lophate perispore with rounded crests (costate to costate-cristate) and very few pori (less than 1 ΅m in diam.), middle perispore present in areolae between outer perispore folds.
* On rocks in forests; 26003600 m. Sichuan.
All specimens of Asplenium tenuicaule var. argutum come from relatively high elevations in central Sichuan. They have slightly longer stipes and relatively long marginal teeth; perispore characters are constant. Due to confusion with similar taxa, the distribution of this taxon is not well known.
Chinese pinyin
Rhizome erect, short, scaly; scales brown to dark brown, 24(5) Χ ca. 0.5 mm. Fronds caespitose, (5)1025(30) Χ (1)3(7) cm; stipe semiterete, (1.2)25(8) cm, adaxially green and sulcate, abaxially entirely green (small fronds) or dark brown for 1/43/4 of its length (in large fronds into lower part of rachis), with scales similar to those on rhizome; lamina ovate, apex acute-acuminate, 2- or 3-pinnate at base; pinnae (6)10(14), subopposite to alternate, with stalks 12 mm, triangular to ovate, 24(5) cm, lowest usually reduced, base asymmetrical, acroscopically truncate, basiscopic side cuneate, apex obtuse to acute; segments 24 pairs, alternate, anadromous, basal acroscopic segment largest and free, others usually adnate to costa, broadly ovate to orbicular. Sori several on each pinna, medial on subtending vein, often confluent at maturity, 1.53 mm. Fronds herbaceous, green; rachis green, sulcate adaxially, abaxially often shiny castaneous at base but green toward apex; lamina with 3- or 4-celled uniseriate gland-tipped hairs, average guard cell length 3740 ΅m. Sori several on each pinna, medial on subtending vein, often confluent at maturity, oval-linear, 1.53 mm; indusia membranous, repand to entire, opening toward costule or to costa. Spores with lophate (cristate-alate) perispore, average exospore length 2427 ΅m. Plant diploid sexual: 2n = 72.
On rocks or trees in forests; 12002500 m. Yunnan [India, Philippines, Sri Lanka; mountains of tropical Africa].
The type is from India.
The range of Asplenium semivarians is not well known due to confusion with similar species of the Asplenium varians complex. It may also occur in Malaysia (Johore) but the single specimen, cited by Morton ([??citation]. 1961) under A. varians, needs further study. Experimental hybridization and micromorphology have shown (Viane & Reichstein, [??citation]. 2003) that it is different from diploid A. tenuicaule and that it is an ancestor of true A. varians. Asplenium semivarians is known from southern India as diploid Asplenium varians (e.g., Manickam & Irudayaraj, [??citation]. 1988; Bir & Irudayaraj, [??citation]. 2001). Luxurious plants become almost tripinnate. This species can be distinguished from similar taxa by its diploid chromosome number, its alate perispore with the middle perispore lacking in the areolae (zones between the ridges), and the smaller exospore and guard cells.
Chinese pinyin
Asplenium atuntzeense Viane & Reichstein; A. daghestanicum Christ subsp. aitchisonii (Fraser-Jenkins & Reichstein) Fraser-Jenkins; A. minutum C. Y. Yang (1993), not Willdenow ex Klotsch (1847); A. subdigitatum Ching (1985), not Mettenius ex Kuhn (1868); A. xinjiangense Ching.
Plants 818 cm tall. Rhizome short, erect to ascending, apex scaly; scales dark brown, ovate-triangular, 24 mm, often with marginal fimbriae. Fronds caespitose, bright to dark green, (2)711(15) cm; stipe usually as long as (up to 1/3 of) lamina, 26 cm, semiterete, base dark brown for 215 mm but green toward rachis, with narrow scales and uniseriate hairs, subglabrous when old; lamina narrowly triangular to ovate, base widest, (2)36 Χ (0.5)12(3) cm, bipinnate at base, apex acute; pinnae 510 pairs, subopposite or alternate, basal pinnae with short stalks 0.52(3) mm, upper pinnae sessile, basal pinnae usually largest, triangular to deltoid, 312(18) Χ 310(15) mm, apex obtuse, lowest pinnae 0.52 cm separated, each usually with two pinnules and broad, deeply pinnatifid apex, upper pinnae smaller, less divided and entire near apex; ultimate segments with cuneiform base and obtuse apex, margin dentate, with 35(7) spreading teeth. Costa obvious but veins obscure, veins anadromously branching. Frond herbaceous, green to brown-green when dried, rachis semiterete, green, with narrow scales and uniseriate hairs; average guard cell length 5661 ΅m. Sori submedian to median on subtending vein, confluent when mature, 23 mm; indusia membranous, repand to entire, opening toward costule or toward costa. Spores with lophate (costate to cristate) perispore, average exospore length 3842 ΅m. Sexual hexaploid species: 2n = 216.
On half-shaded, non-calcareous rocks; 17004200 m. S Gansu, Sichuan, Xinjiang, Xizang, Yunnan [Bhutan, N India, Nepal, Pakistan].
Asplenium aitchisonii is similar to other members of the Asplenium varians-complex (i.e., A. altajense, A. kukkonenii, A. semivarians, A. tenuicaule, A. varians), but is usually widest at the base and has longer stomates and spores due to its higher number of chromosomes. This is an allohexaploid (not an octoploid as stated in the original description). Asplenium daghestanicum, a tetraploid from the Caucasus and Libya (Tibesti), is genetically isolated from this species and thus better kept as a separate species. Plants from isolated populations, growing in sheltered conditions in regions with an extreme climate, are often less developed and have been described as separate species (i.e., A. xinjiangense, A. subdigitatum, A. minutum).
朗木铁角蕨 lang mu tie jiao jue
Plants 58 cm tall. Rhizome erect, short, apex scaly; scales dark brown, narrowly triangular, subentire to denticulate. Fronds caespitose; stipe 1.53.5 cm, slender, adaxially sulcate, base dark brown and with scales, upward green, with fibrillar scales; lamina narrowly triangular, 45 Χ 1.41.8 cm, base slightly reduced, apex acute-acuminate with triangular apical segment ca. 12 Χ 6 mm, 2-pinnate at base; pinnae 5 or 6 pairs, opposite to subopposite, shortly stalked, basal pinnae slightly reduced, middle pinnae trapeziform-elliptic, 69 Χ 49 mm, base subsymmetrical truncate to cuneate, 1-pinnate at base or simple, apex obtuse; basal pinnae with up to 2 pinnule pairs, basal acroscopic pinnules largest and free, 34 Χ 22.5 mm, base broadly cuneate, apex serrate with long and sharp teeth, apex obtuse. Veins slightly raised adaxially, 2-forked or simple, not reaching margin. Fronds firmly herbaceous, green, brown-green when dry; rachis green, adaxially sulcate, lamina subglabrous, average guard cell length 4552 ΅m. Sori 13 per segment, median on subtending vein, confluent at maturity, linear, 23 mm; indusia white-gray, semi-elliptic, membranous, entire, opening toward costa or costule, persistent. Spores with lophate (costate) perispore, average exospore length 3336 ΅m.
* Juniperus forests; 33003600 m. S Gansu, NW Sichuan.
The type is from Sichuan (S of Langmu Si).
Asplenium neovarians, known from few collections, needs more study. Based on the length of its stomates and exospore, it should at least be tetraploid. It is similar to A. altajense (but different in its more distinct apical pinna), and to A. aitchisonii (but different in its putative ploidy level), and to A. kukkonenii, from all of which it differs in perispore morphology.
Chinese pinyin
Type: China. Inner Mongolia (Nei Menggu): Hulunbuir, Balin, Beishan, in rock crevices, 460 m, 2 Aug 1983, Y.-Q. Ma et al. 83-7 (holotype, HIMC).
Planta morphologia Asplenio tenuicaulis. Rhizoma breve adscendens vel erectum, squamis 1.52 mm longiset ca. 0.5 mm latis. Folia caespitosa, maturitate (4)615(20) cm longa, basi 13 cm lata. Petiolus lamina 2 plo brevior, pro parte abaxiale basale Ό ad Ύ fuscus, basi squamis rhizomatis similibus. Lamina anguste triangularis-ovata, basi bipinnata, paribus pinnarum suboppositarum vel alternantium 510. Jugum primum vel secundum maximum, pinnis (5)812(20) mm longis. Pinnae triangulares-ovatae petiolulis ad 1 mm, inaequilaterae petiolum versus truncatae, basin cuneatae, apice obtusae. Lamina papyracea, stomatibus 4953 ΅m longibus. Sori ca. 0.82.5 mm longi, maturitate confluentes. Sporae exosporium 3439 ΅m longum, perisporium costatum.
Plants (4)615(20) cm tall. Rhizome erect, short, apex scaly; scales brown to dark brown, triangular to narrowly triangular, 1.52 Χ 0.40.5 mm, base hastate-cordate, margin glandular, fimbriate or entire. Fronds caespitose; stipe green but base castaneous, brown color occasionally extending into rachis, (1)1.55.5(7) cm, with small dark brown, fimbriate scales and uniseriate hairs, subglabrous when old, adaxially sulcate; lamina triangular to narrowly triangular-ovate, (2.5)3.56.5(9) Χ 13 cm, base truncate to broadly cuneate, pinnate-pinnatifid to bipinnate, apex acute-acuminate; pinnae 510 pairs, subopposite to alternate, stalk up to ca. 1 mm, basal or first suprabasal pinnae pair largest, triangular-ovate, (5)812(20) Χ (3)57(8) mm, base asymmetrical, acroscopic side truncate, basiscopic side cuneate, pinnate, rarely 2-pinnatifid, apex obtuse; pinnules 2 or 3 pairs, anadromous, basal pinnules free or adnate to costa, margin serrate with acute to mucronate teeth, apex obtuse. Veins obscure, veinlets 1- or 2-forked, not reaching margin. Fronds thinly herbaceous, green to gray-green or brown when dried, rachis green, only rarely with castaneous base (abaxially), adaxially sulcate, subglabrous, lamina with 3- or 4-celled uniseriate gland-tipped hairs, average guard cell length 4953 ΅m. Sori 13 per pinnule, median on acroscopic veinlets, occasionally confluent at maturity, oval-linear, 0.82.5 mm; indusia whitish to pale brown, oval-linear, membranous, entire to repand, opening toward costule or costa. Spores with lophate (costate-cristate) perispore, average exospore length 3439 ΅m.
* In rock crevices; 4001800 m. Liaoning, Nei Mongol (Da Xingan Ling), Shanxi.
This species is named after Professor Yu-Quan Ma, principal redactor of Flora Intramongolica.
Asplenium mae is mainly known from the Greater Hinggan Range. It is a putative autotetraploid which probably originated via chromosome doubling in A. tenuicaule var. subvarians, which it strongly resembles. Average spore size indicates that it is probably tetraploid, but no chromosome counts are yet available. In herbaria, A. mae is either filed under A. tenuicaule, A. subvarians, A. anogrammoides, or A. sarelii.
Chinese pinyin
Plants (3.5)615(25) cm tall. Rhizome erect, short, apex scaly; scales brown to dark brown, triangular to narrowly triangular, (2)2.23.5(4.5) Χ 0.40.7 mm, fimbriate or entire. Fronds caespitose; stipe green (small fronds) but base dark brown, color occasionally extending into rachis, (1)26(9) cm, with small dark brown, often linear, scales and uniseriate hairs, subglabrous when old, adaxially sulcate; lamina narrowly oblong to ovate, (2.3)510(20) Χ (0.7)12.5(4) cm, base truncate-cuneate, bipinnate at base, pinnate-pinnatisect (-pinnatifid) in upper part, apex acute-acuminate to caudate; pinnae 713 pairs, alternate or opposite, stalk ca. 1 mm, lower pinnae slightly reduced, lowest pair 0.81.8 cm separated from next, more distal pinnae closer but usually not overlapping, median and upper pinnae often at an angle of 4575° to rachis, largest pinnae triangular-ovate, 1020 Χ 510(12) mm, base asymmetrical, acroscopic side truncate, basiscopic side cuneate, basal acroscopic pinnule largest, often shortly (ca. 0.5 mm) stipitate and nearly parallel to rachis, fan-shaped or ovate to triangular-ovate, margin serrate to crenate with acute to mucronate teeth, apex obtuse; basiscopic pinnules often adnate and directed outward, almost perpendicular to rachis. Fronds herbaceous, green to gray-green or brown when dried, rachis green or occasionally with castaneous base (abaxially), adaxially sulcate, with small, dark brown, linear scales, lamina with 3- or 4-celled uniseriate gland-tipped hairs, epidermis cells with sinuous pseudomamillate anticlinal walls, average guard cell length 4550 ΅m. Sori 13(or 4) per pinnule, median on acroscopic veinlets, often confluent at maturity, oval-linear, 1.42.4(3) mm; indusia whitish to pale brown, oval-linear, membranous, entire to repand, opening toward major veins or costa. Spores with lophate (costate-cristate) perispore, average exospore length 3236 ΅m. Plants sexual, allotetraploid: 2n = 144.
On rocks in forests; 21003300 m. Xizang, Yunnan [India, Nepal, Pakistan].
Asplenium kukkoneii is most similar to A. tenuicaule and A. varians. Crossing experiments and micromorphology have shown that A. tenuicaule is one of its ancestors (Viane & Reichstein, [??citation]. 2003). The second ancestor of A. kukkonenii is still unknown. The fact that A. kukkonenii and A. varians share a genome explains why these taxa are difficult to differentiate from each other. The lamina of A. kukkonenii is usually a little narrower and more acuminate (occasionally even caudate) at the apex. The mean length of the rhizome scales is usually less than 3.5 mm in A. kukkonenii, and more than 3.5 mm in A. varians. Reliable differentiation is also possible by examination of epidermal cells, which are thickened or pseudomamillate in their sinuses in A. kukkonenii, but not so in A. varians. In herbaria, A. kukkonenii is often identified as A. varians or even A. fontanum.
Chinese pinyin
Asplenium fimbriatum Kunze var. leptophyllum Zenker ex Kunze; A. propinquum Ching; A. saulii Baker var. latius Christ, p.p.
Plants 515(25) cm tall. Rhizome erect, short, apex scaly; scales dark brown, narrowly triangular, 23.5 mm, fimbriate at base or entire. Fronds caespitose; stipe semiterete, 28(10) cm, adaxially green and sulcate, abaxially shiny castaneous, brown color occasionally extending into rachis, with small dark brown, fimbriate scales and uniseriate hairs, subglabrous when old; lamina triangular-ovate, (2)512(15) Χ 13(6) cm, apex acute-acuminate, 2- or 3-pinnate; pinnae 812 pairs, alternate or basal ones opposite, stalks 12 mm, lower pinnae usually somewhat reduced, middle pinnae triangular-ovate, 817(25) Χ 712 mm, base asymmetrical, acroscopically truncate, basiscopic side cuneate, 1- or 2-pinnate, apex obtuse; pinnules 24 pairs, alternate, anadromous, basal acroscopic pinnule largest and most divided, broadly ovate, 68 Χ 46 mm, base broadly cuneate, shortly stalked, pinnatifid or pinnate, apex obtuse; other pinnules oval to cuneiform, 3 or 4 pairs, reduced toward pinna apex, apical margin serrate and with long, sharp teeth. Costa obvious but veins obscure, veins anadromously branching, often with terminal submarginal hydathode. Fronds thinly herbaceous, when dry grass-green; rachis same color as stipe and sulcate adaxially, abaxially often shiny castaneous at base, becoming green toward apex; lamina with 3- or 4-celled uniseriate gland-tipped hairs, average guard cell length 4652 ΅m. Sori 13 per pinnule, submedian to median on subtending vein, shortly linear, 13 mm; indusia membranous, repand to entire, opening toward costule or costa. Spores with reticulate perispore, average exospore length 3337 ΅m.
On rocks in forests; 11002000 m. Hebei, Hubei, Shaanxi, Shanxi, Sichuan, Yunnan [S India].
The type is from Shaanxi.
Asplenium kansuense is a putative autotetraploid, which probably originated via chromosome doubling in A. tenuicaule var. tenuicaule. Its general morphology and perispore are very similar to those of that taxon, but its spore size indicates that it is probably tetraploid. No chromosome counts are available. In China, A. kansuense was often misidentified as A. varians or as A. sarelii sensu lato. Large tripinnate plants resemble tetraploid A. anogrammoides with which they share half of their genome.
The southern Indian (Nilgiri) plants belonging to this species (Asplenium fimbriatum var. leptophyllum Kunze) occur at higher elevations, are smaller than the Chinese plants, and are similar to both A. varians and A. semivarians. Several chromosome reports of S Indian tetraploid Asplenium varians may refer to this species. Large specimens of African A. fimbriatum Kunze are closely similar, also in perispore characters. Hybridization experiments have shown that true A. fimbriatum is an allotetraploid with the same genome composition as A. varians and is thus best considered a subspecies or a variety of the latter (Viane & Reichstein, [??citation]. 2003).
变异铁角蕨 bian yi tie jiao jue
Asplenium laciniatum D. Don, nom. rej. (ICBN Art. 57.1), not A. laciniatum sensu Hooker et auct. plur. = A. gueinzianum Mettenius ex Kuhn; A. lankongense Ching; A. paucijugum Ching (1985), not F. Ballard (1935).
Plants (4)820(35) cm tall. Rhizome erect, short, apex scaly; scales dark brown to black, narrowly triangular, (1.5)34.2(5) Χ 0.51 mm, fimbriate at base or subentire. Fronds caespitose; stipe semiterete, (0.5)27(12) cm, adaxially green and sulcate, abaxially shiny castaneous, brown color occasionally extending to rachis, with small dark brown, very narrow scales and uniseriate hairs, subglabrous when old; lamina triangular-ovate, (3)613(25) Χ (1)24(8) cm, base slightly reduced, bipinnate, apex acute-acuminate; pinnae (5)812(18) pairs, lower pinnae (sub)opposite, upper alternate, stalk ca. 1 mm, largest pinnae triangular-ovate, 817(30) Χ (3)610(15) mm, base asymmetrical, acroscopically semicordate-truncate, basiscopic side cuneate, 1-pinnate, apex obtuse; pinnules 2 or 3 pairs, alternate, anadromous, basal acroscopic pinnule largest, fan-shaped to orbicular-obovate, 3.55.5 Χ 2.54(6) mm, base cuneate, sessile to almost free, lateral side entire, apex serrate and obtuse; other segments smaller, usually adnate to costa. Veins not obvious, slightly raised adaxially when dry, anadromously forking, not reaching margin. Frond herbaceous, green or grayish green when dry; lamina with 3- or 4-celled uniseriate gland-tipped hairs, average guard cell length 4652 ΅m; rachis semiterete, grayish green and sulcate adaxially, occasionally brown at base abaxially, subglabrous. Sori 24 per pinnule, submedian to median on subtending vein, often confluent at maturity, oval to linear, (1)1.53(4.5) mm; indusia gray, linear, membranous, repand to entire, opening toward costule or costa, persistent. Spores with cristate-alate perispore, average exospore length 3135 ΅m. Plants sexual allotetraploid: 2n = 144.
On wet rocks or cliffs in forests, or on man-made constructions[??buildings, walls]; 5004200 m. Chongqing, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Shaanxi, Sichuan, Xizang, Yunnan [Bhutan, India, Nepal, Vietnam; S Africa].
Sleep and Reichstein ([??citation]. 1984) discussed the problem of the application of the name Asplenium laciniatum D. Don (1825) in their typification of A. varians Wallich ex Hooker & Greville (1830), which is a later synonym of A. laciniatum. The problem with D. Dons name is caused by Moore (1857) and Hooker (1860), who applied the epithet to another Himalayan fern not related to A. varians. Most later authors followed their concept of A. laciniatum (e.g., Beddome, [??citation]. 1864; [??citation]. 1892; Hooker & Baker, [??citation]. 1867; [??citation]. 1874; Clarke, [??citation]. 1880; Franchet, [??citation]. 1885; Blanford, [??citation]. 1888; Diels, [??citation]. 1902; Christ, [??citation]. 1905; Christensen, [??citation]. 1905; Hayata, [??citation]. 1914; Hieronymus, [??citation]. 1919; Blatter & dAlmeida, [??citation]. 1922; Ogata, [??citation]. 1930; Wu et al., [??citation]. 1932; Mehra & Bir, [??citation]. 1964; Ito, [??citation]. 1966; [??citation]. 1971; Nishida et al., [??citation]. 1966; Tagawa et al., [??citation]. 1966; Bir, [??citation]. 1968; [??citation]. 1973; [??citation]. 1987; Subramanyam, [??citation]. 1973; Nayar & Kaur, [??citation]. 1974; De Vol et al., [??citation]. 1975; Iwatsuki, [??citation]. 1975; [??citation]. 1988; Malla et al., [??citation]. 1976; Dhir, [??citation]. 1980; Chandra & Kaur, [??citation]. 1984; Dixit, [??citation]. 1984; [??citation]. 1989; Baishya & Rao, [??citation]. 1982; Iwatsuki et al., [??citation]. 1986; Jamir & Rao, [??citation]. 1988; Zhang et al., [??citation]. 1990; Khullar, [??citation]. 1992; Ni & Cheng, [??citation]. 1992; Pande & Pande, [??citation]. 1993; Shieh et al., [??citation]. 1994; Wu, [??citation]. 1999; and G.F. Zhang, [??citation]. 2006)[??shorten list of examples??].
Because the name Asplenium laciniatum D. Don has thus been widely and persistently used for a taxon not including its type, for nomenclatural stability (ICBN Art. 57.1), this name should be rejected. This rejection has the advantageous side effect of preserving the present usage of the name A. varians Wallich ex Hooker & Greville. Whether A. laciniatum D. Don is rejected or not, the correct name for the taxon with the misapplied name is A. gueinzianum (Morton, [??citation]. 1973; Viane & Reichstein, [??citation]. 1986; Khullar, [??citation]. 1994).
This species is variable, very similar to, and often confused with its ancestors, Asplenium semivarians and A. tenuicaule (see above). Its relationships were discovered using hybridization experiments, chromosome counts, and micromorphology (Sleep & Reichstein, [??citation]. 1984; Viane & Reichstein, [??citation]. 2003), showing it is an allotetraploid species that arose by chromosome doubling in the diploid hybrid between A. tenuicaule (var. subvarians in A. varians var. varians, and var. tenuicaule in the African var. fimbriatum) and A. semivarians. Two varieties are distinguished: A. varians var. varians occurring in the Himalaya from northern India into China (not in Japan), and var. fimbriatum, which is mainly found in southern Africa (though its occurrence in southern India needs further study). Asplenium paucijugum Ching is similar to A. semivarians but has larger spores and stomates. Further study is needed to show if this and related forms are autotetraploids of A. semivarians.
阿尔泰铁角蕨 a er tai tie jiao jue
Asplenium sarelii Hooker [saulii] var.[??f.] altajense Komarov, Bull. Jard. Bot. Petrop. 16: 150. 1916; A. chingianum Chang Y. Yang.
Plants 515 cm tall. Rhizome erect, short, apex scaly; scales dark brown to black, narrowly triangular, 23.5 Χ 0.30.5 mm, entire. Fronds caespitose; stipe 2.53.5(6) cm, base scaly and brown abaxially, green upward, adaxially green, sulcate, subglabrous; lamina 2-pinnatifid to 2-pinnate, narrowly triangular-ovate, 89 Χ 22.5 cm, base slightly reduced, apex acute; pinnae (5)1015 pairs, lower pinnae subopposite, often deltoid-flabellate or almost orbicular, upper alternate and gradually fused into 0.51 cm pinnatifid apex, second to fourth pair usually largest, stalk up to 1 mm and adaxially sulcate with raised supravascular ridge, middle pinnae triangular-ovate, ca. 11 Χ 810 mm, base asymmetrical, acroscopic side truncate, basiscopic side cuneate, 1-pinnate, apex obtuse; pinnules 2 or 3 pairs, anadromous, basal acroscopic pinnule free, largest, subovate, 45 Χ 24 mm, base cuneate, other pinnules adnate to costa and confluent into pinna apex, margins dentate-serrate, apex obtuse. Veins obscure, flabellately anadromous, slightly raised adaxially. Fronds papery, green to grayish green when dry, lamina subglabrous, average stomatal guard cell length 4448 ΅m; rachis semiterete, green, adaxially sulcate and with raised supravascular ridge. Sori 24 per pinnule, basal to medial on pinnae, at maturity confluent, linear-elliptic, 12.5 mm; indusia grayish green, linear-elliptic, membranous, entire, opening toward costa or costules. Spores with lophate (cristate) perispore, average exospore length 3234 ΅m. Plants sexual, allotetraploid: 2n = 144.
In crevices of shaded rocks; 4001500 m. Nei Mongol, Ningxia, Sichuan, Xinjiang [Mongolia, Russia (Altai, S Siberia)].
Asplenium altajense is relatively common in the area bordering China, the Altai Mountains, Mongolia, and Russia, but future research will have to show if the area given by Grubov ([??citation]. 1963) is correct. In general shape, this taxon is similar to A. pekinense Hance. However, most pinnae of A. altajense have only the basal acroscopic pinnule free, the others are confluent and form the pinna apex, while in A. pekinense several segments are free or just slightly adnate to the costa. The stipe is less scaly in A. altajense, and its texture is more herbaceous. Asplenium altajense is also similar to A. tenuicaule, from which it can be distinguished by the more narrow and acute teeth (obtuse to mucronate in A. tenuicaule). Due to confusion between A. tenuicaule, A. pekinense, and A. anogrammoides, the distribution of this taxon is not well known. Asplenium chingianum (type fragment in IBSC!) is a submature plant with immature sporangia, few and unripe spores, and almost no scales; it is also similar to A. nesii specimens from N Pakistan. The hybrid between A. altajense and A. tenuicaule is relatively common where they grow together: Asplenium Χshmakovii Viane, hybr. nov. Type: Russia. Altai Republic: ca. 35 km S-SSE of Chemal, N exposed limestone-marble rocks in R. Karasu valley, W of R. Katun, 580 m alt., 6 Aug 2004, Viane 10000A (holotype, GENT). Planta hybrida, inter parentes A. altajensem et A. tenuicaulem intermedia, sporis abortivis, numero chromosomatico triploideo 2n = 108, meiosi 36 bivalentibuset 36 univalentibus differt. Meiotic chromosome behavior in this morphologically intermediate hybrid shows that A. altajense contains one chromosome set of A. tenuicaule.
黑色铁角蕨 hei se tie jiao jue
Asplenium adiantum-nigrum subsp. yuanum (Ching) Reichstein et al.; A. adiantum-nigrum var. yuanum (Ching) Ching; A. yuanum Ching.
Plants 1540 cm tall. Rhizome ascending, apex scaly; scales dark brown, narrowly triangular, up to 6 mm, hairlike in upper half, without black middle stripe, entire. Fronds caespitose; stipe 815(20) cm, base conspicuously swollen, thickened, semiterete, abaxially shiny castaneous to blackish purple, with scattered brownish black hairlike scales or subglabrous, adaxially sulcate; lamina triangular-ovate, 925 Χ 46(10) cm, apex acute to acuminate, tripinnate; pinnae 813 pairs, alternate, stalked, basal pair largest, triangular, 46 Χ 1.82.4 cm, base truncate, bipinnate, apex acute to acuminate; pinnules 58 pairs, anadromous, basal pairs largest, triangular, 1.53.2 Χ 0.61 cm, base cuneate-truncate, shortly stalked or subsessile, pinnate, apex acute; ultimate pinnules 46 pairs, alternate, anadromous, basal pair largest, elliptic to oblong or linear, 57 Χ 2.53.5 mm, margin serrate, apex obtuse. Costa obvious, sulcate adaxially, veins obscure, anadromously branching, simple or 2-forked, not reaching margin. Fronds firmly herbaceous to papery, green but brown-green when dry, lamina with 4- or 5-celled, uniseriate hairs, average stomatal guard cell length 5460 ΅m; rachis semiterete, lower half abaxially usually shiny castaneous to dark brown, apical part green or stramineous, adaxially sulcate. Sori 13(or 4) pairs per pinnule or segment, median on subtending veinlet, confluent at maturity, linear, 12.5(5) mm; indusia white to brownish, linear, membranous, repand to entire, opening toward costa or costule. Spores brown to dark brown, perispore lophate (cristate-alate), with average exospore length 3237 ΅m. Plants sexual allotetraploids: 2n = 144.
On rocks or on forest floor near streams; 20002800 m. Shaanxi, Taiwan, E Xizang, NW Yunnan [Afghanistan, N India, Pakistan; Europe (Mediterranean to W Asia), W North America].
The type is from Europe.
Asplenium adiantum-nigrum, an allotetraploid species that originated via chromosome doubling in a hybrid (A. Χligusticum) between A. cuneifolium Viviani and A. onopteris, is unrelated to distantly similar A. interjectum. Asplenium adiantum-nigrum differs by the thickened stipe base (trophopod) and the less deltoid lamina.
Chinese pinyin
Asplenium leiboense Ching; A. sarelii Hooker var. anogrammoides (Christ) Tagawa; A. sepulchrale Hooker, p.p.; A. wudangense Z. R. Wang & X. Hou[IPNI: nom. inval. without Latin description (check Acta Bot. Sin. 45: 4. 2003)].
Plants 1030 cm tall. Rhizome short, erect or ascending, apex scaly; scales dark brown to black, narrowly triangular, 38 Χ 0.30.7 mm, margins denticulate; scale base hyaline, cordate, with numerous yellow-brown, unicellular hairs 15 Χ 0.02 mm, similar to root hairs. Fronds caespitose; stipe 715 cm, semiterete, dark brown abaxially or becoming green toward rachis, upward green, adaxially green, shallowly sulcate with supravascular ridge, with similar scales as on rhizome, reduced hairlike scales toward rachis or subglabrous; lamina ovate, 613 Χ 2.57(8) cm, base truncate, apex acute, tripinnatifid-tripinnate; pinnae 812 pairs, basal pairs subopposite to alternate, stalk 24 mm, sulcate adaxially; several basal pinnae pairs about equal in length, ovate-triangular, 1.53 Χ 0.51.5 cm, base subsymmetrical, truncate to broadly cuneate, 2-pinnatifid-bipinnate, apex acute; pinnules 46 pairs, anadromous, acroscopic and basiscopic pinnules about equal in size, narrowly triangular to ovate, 510 Χ 36 mm, base asymmetrical, acroscopic side truncate, basiscopic side cuneate, largest pinnules stalked, others decurrent on costa, pinnatipartite-pinnate, apex acute; segments 2 or 3 pairs, linear to narrow elliptic, apex with 2 or 3 teeth. Costa sulcate adaxially, without median ridge, veins slightly raised adaxially, anadromously forked. Fronds firmly herbaceous, green to grayish green, lamina subglabrous, average stomatal guard cell length 4852 ΅m; rachis green, semiterete, abaxially green or base castaneous becoming green toward apex, adaxially sulcate with raised median supravascular ridge, with reduced scales or subglabrous. Sori 24 per ultimate segment, median to subterminal on acroscopic vein, subconfluent at maturity, subelliptic, 12.5 mm; indusia grayish green, subelliptic, membranous, entire, opening toward costule or central veinlet, persistent. Spores brown, perispore lophate (cristate), average exospore length 3336 ΅m. Plants sexual, allotetraploids: 2n = 144.
Wet cliffs, in crevices of limestone rocks, on man-made constructions[??buildings, walls]; 3002800 m. Anhui, Fujian, Guangdong, Guizhou, Hebei, Hubei, Hunan, Jiangsu, Jiangxi, Jilin, Liaoning, Ningxia, Shaanxi, Shandong, Shanxi, Sichuan, Yunnan, Zhejiang [?India, Japan, Korea, Vietnam].
The type is from Korea.
Asplenium anogrammoides is an (allo)tetraploid (Kurita, [??citation]. 1960; Mitui, [??citation]. 1968) and originated via chromosome doubling in the sterile hybrid between A. sarelii and A. tenuicaule (Lin & Sleep, [??citation]. 1989; Murakami et al., [??citation]. 1999; Viane & Reichstein, [??citation]. 2003; Wang, [??citation]. 2003). Their morphological similarity led to confusion and misapplication of the name A. sarelii to this taxon (tetraploid sarelii). Moreover less divided plants of A. anogrammoides are similar to large plants of A. pekinense (the autotetraploid arisen from A. sarelii). On the other hand, the name A. anogrammoides has been misapplied (e.g., Fomin, [??citation]. 1930; [??citation]. 1934) to southern Siberian and Mongolian A. tenuicaule var. subvarians.
Yellow-brown (root)hairs are present on the rhizome, stipe base, and base of the scales of A. pekinense, A. anogrammoides, A. altajense, A. sarelii, A. tenuicaule, and other related taxa, and thus cannot be used to distinguish them. The average exospore and stomata length are the most reliable characters discriminating between diploid and tetraploids in this complex. Asplenium anogrammoides can often be separated from A. pekinense, by its largely unreduced lamina base, and its ovate-triangular pinnae (deltoid in A. pekinense) with stalks 24 mm (0.52 mm in A. pekinense).
Where Asplenium anogrammoides grows together with A. tenuicaule, their sterile triploid hybrid, A. Χmitsutae, is common. Asplenium Χmitsutae Viane & Reichstein, hybr. nov. Type: Japan. Kyoto pref.: Kibune shrine, ca. 12 km N of Kyoto city, on stone wall along road, half shaded limestone area, ca. 200 m alt., 1 Dec. 1985, S. Mitsuta, H. Iwahori & Y. Hosoi s.n. (TR-6350) (holotype, G). Planta hybrida, inter parentes A. anogrammoideum et A. tenuicaulem intermedia, sporis abortivis, numero chromosomatico triploideo 2n = 108, meiosi 36 bivalentibus 36 univalentibusque differt. Meiotic chromosome behavior in this hybrid shows that A. anogrammoides contains one chromosome set of A. tenuicaule. This hybrid is named after S. Mitsuta, who collected several living plants for T. Reichsteins research. Asplenium sarelii var. magnum H. S. Kung. described from Sichuan, is probably the same hybrid combination.
华中铁角蕨 hua zhong tie jiao jue
Asplenium blakistonii Baker; A. pekinense Hance var. foeniculaceum Christ; A. saulii Hooker ex Baker[saulii was a misspelling/interpretation of the handwritten name sarelii].
Plants 1023 cm tall. Rhizome short, erect or ascending, apex scaly; scales dark brown to black, narrowly triangular, 24(6) Χ 0.30.7 mm, margins denticulate; scale base hyaline, cordate, with yellow-brown, unicellular hairs 15 Χ ca. 0.02 mm, similar to root hairs. Fronds caespitose; stipe 710 cm, semiterete, base dark brown abaxially, upward green, adaxially yellow-green, sulcate with pronounced supravascular ridge, with similar scales as on rhizome, reduced hairlike scales toward rachis or subglabrous; lamina triangular-ovate, 615 Χ 2.57 cm, base truncate, apex acute, tripinnatifid-tripinnate; pinnae 812 pairs, basal pairs subopposite to alternate, stalk 13 mm, sulcate but with median supravascular ridge adaxially; basal pinnae equal or slightly shorter than next, ovate-triangular, 1.54.5 Χ 1.53 cm, base subsymmetrical, truncate to broadly cuneate, bipinnatifid-bipinnate, apex acute; pinnules 46 pairs, anadromous, basal pinnules about equal in size, narrowly triangular to ovate, 520 Χ 47 mm, base asymmetrical, acroscopic side truncate, basiscopic side cuneate, stalk up to 1 mm or decurrent on costa, pinnatipartite-pinnate, apex acute; segments 3 or 4 pairs, linear, 1.55 Χ 0.52 mm, basal segments forked or up to pinnatisect with 2 or 3 ultimate segments, apex of segments with 2 or 3 teeth. Costa sulcate adaxially, with distinctly raised median supravascular ridge, veins raised adaxially, veins anadromously forked. Fronds thinly to firmly herbaceous when dry, green to grayish green, lamina subglabrous, average stomatal guard cell length 3543 ΅m; rachis green, semiterete, adaxially sulcate with raised median supravascular ridge, with reduced scales or subglabrous. Sori 1 or 2 per ultimate segment, median to subterminal on acroscopic vein, near segment teeth but not reaching margin, subconfluent at maturity, subelliptic, 12 mm; indusia grayish green, subelliptic, membranous, entire, opening toward costule or central veinlet, persistent. Spores brown, perispore lophate (cristate), average exospore length 2830 ΅m. Plants sexual diploids: 2n = 72*.
* In rock crevices; 3001000(2100) m. Anhui, Chongqing, N Guizhou, ?Henan, Hubei, Hunan, Jiangsu, Shaanxi, Sichuan, Zhejiang.
The type is from Chongqing or Sichuan.
Asplenium sarelii is endemic to central China. All reports of this species from outside China are erroneous and usually refer to the very similar tetraploid A. anogrammoides. Hybridization experiments, including cytological study of meiotic behavior in artificially produced hybrids (Lin & Sleep, [??citation]. 1989), and micromporphological study of type material have shown that A. sarelii is one of the parental species of allotetraploid A. anogrammoides; the other parent being diploid A. tenuicaule. These findings were confirmed by cytological and isozyme studies (Wang, [??citation]. 2003). The morphological similarity between these taxa has led to great confusion, also with true A. pekinense, which is the autotetraploid that originated from true A. sarelii. In A. sarelii, the ultimate segments are less than 1.2 mm wide, the pinnae stalks less than 1 mm wide, and its pinnae are distant from each other and hardly overlap so that the frond silhouette is more open than in A. anogrammoides. Important differential characters are the chromosome number and the average exospore and stomata size. Identification based on morphology alone is very difficult when A. sarelii grows with A. pekinense and A. anogrammoides. At such locations, e.g., Wudang Shan (Hubei) and Huping Shan (Hunan), the sterile triploid hybrids, A. Χwudangshanense (A. pekinense Χ A. sarelii) and A. Χhuawuense (A. anogrammoides Χ A. sarelii), are relatively common and recognizable by their aborted spores and often undeveloped sporangia. Asplenium Χwudangshanense Viane, Reichstein, Rasbach & Y.X. Lin, hybr. nov. Type: China. Hubei: Wudang Shan, ca. 2050 m alt., Sep 1907, Silvestri 6 (pl. 5) (holotype, FI). Planta hybrida, inter parentes A. pekinensem et A. sarelium intermedia, sporis abortivis, numero chromosomatico triploideo 2n = 108, meiosi 05 trivalentibus, 3337 bivalentibus 2940 univalentibusque differt. Meiotic chromosome behavior in A. Χwudangshanense confirms that A. pekinense is the autotetraploid of A. sarelii. Meiotic chromosome behavior (Wang, [??citation]. 1989; [??citation]. 2003) in A. Χhuawuense confirms that A. anogrammoides is allotetraploid and contains one chromosome set of A. sarelii. Asplenium Χhuawuense Z. R. Wang ex Viane & Y. X. Lin, hybr. nov. Type: China. Hubei, Wudang Shan, ca. 850 m, 18 Jul 1987, Z. R. Wang C843 (holotype, PE). Planta hybrida, inter parentes A. anogrammoideum et A. sarelium intermedia, sporis abortivis, numero chromosomatico triploideo 2n = 108, meiosi 36 bivalentibus 36 univalentibusque differt. All of Z. R. Wangs new names are nomina nuda (Wang, [??citation]. 2003).
北京铁角蕨 bei jing tie jiao jue
Asplenium abbreviatum Makino; A. sarelii Hooker subsp. pekinense (Hance) Fraser-Jenkins, Pangtey & Khullar; A. sarelii var. pekinense (Hance) C. Christensen; A. sepulchrale Hooker, nom. illeg.[check (Syn. Fil. ed. 1: 213. 1867); p.p. under sp. 80][Blakiston s.n (type of Asplenium sarelii Hook.) is among the six syntypes, the others are either A. pekinense or A. anogrammoides].
Plants 320 cm tall. Rhizome erect, short, apex scaly; scales dark brown to black, triangular, 15(8) Χ 0.30.7 mm, scale base with numerous yellow-brown, unicellular hairs 15 Χ ca. 0.02 mm, similar to root hairs, apical part subentire, acuminate. Fronds caespitose; stipe green, 28(10) cm, semiterete to terete, base densely scaly, with reduced hairlike scales toward rachis or subglabrous; adaxially sulcate with pronounced supravascular ridge; lamina lanceolate to narrowly ovate-trullate, 515 Χ 1.54 cm, base often gradually reduced, 2-pinnate to 3-pinnatifid, apex acute; pinnae 612 pairs, lower pinnae often reduced, 12.5 cm, stalk 0.51.5(2) mm, adaxially sulcate and with distinct supravascular ridge; basal pinnae remote, opposite, often flabellate to deltoid-triangular, apex obtuse to acute, base symmetrical, truncate, 2-pinnate, with 24 pinnule pairs, basal pinnules nearly parallel to rachis; middle pinnae often largest, triangular-ovate, base asymmetrical, pinnate to bipinnatifid with 24 pairs of pinnules, basal pinnules largest, 58 Χ 24 mm, pinnatisect with 1 or 2 pairs of narrowly cuneiform to sublinear segments, 0.52 mm wide, apex subacute to truncate with 2 or 3 acute, 0.51 mm teeth. Costa sulcate adaxially, with distinctly raised median supravascular ridge, veins obvious, raised adaxially, flabellately anadromous, not reaching margin. Fronds firmly herbaceous, dark green to grayish green when dry, lamina subglabrous, average stomatal guard cell length 4552 ΅m; rachis and costa green, semiterete, adaxially sulcate and with prominent median supravascular ridge, with black fibrillar scales or subglabrous. Sori 1 or 2(4) per pinnule, often spreading at maturity, subelliptic to linear, 12 mm; indusia white-gray, subelliptic, membranous, margin irregularly sinuate, opening toward costa or costule, persistent. Spores brown with lophate (costate) perispore, average exospore length 3438 mm. Plants autotetraploid: 2n = 144.
In crevices of rocks, on man-made constructions[??buildings, walls]; 4003900 m. Anhui, Chongqing, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hebei, Henan, Hubei, Hunan, Jiangsu, Liaoning, Nei Mongol, Ningxia, Shaanxi, Shandong, Shanxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [India, Japan, Korea, Pakistan, Russia (E Siberia)].
The type is from China (Beijing).
Asplenium pekinense is an autotetraploid (Mitui, [??citation]. 1965; Lin & Sleep, [??citation]. 1989; Wang, [??citation]. 2003) species arisen by chromosome doubling in diploid A. sarelii. Allotetraploid A. anogrammoides is very similar because true A. sarelii is one of its ancestors, the second being A. tenuicaule (Lin & Sleep, [??citation]. 1989). Reticulate hybridizations between diploid A. tenuicaule, A. sarelii, and A. semivarians led to the evolution of many confusingly similar taxa defying description. These can be grouped into the A. pekinense-complex consisting of A. altajense, A. anogrammoides, A. pekinense, and A. sarelii, and the A. varians group containing A. aitchisonii, A. kukkonenii, A. semivarians, and A. varians. Their close relationship and similarity has led to many wrong identifications, as well as incorrect and unreliable citations. In many places where A. pekinense grows with A. anogrammoides, their sterile tetraploid hybrid, A. Χkidoi, is relatively common. This morphologically intermediate plant often shows hybrid vigor and was first studied by A. Sleep (in Lin & Sleep, [??citation]. 1989). The name that she proposed is validated here: Asplenium Χkidoi Sleep ex Viane, Y. X. Lin & Reichstein, hybr. nov. Type: Japan. Kyushu: Kumamoto Pref., Ashi-Kita-gun, Usuguchi, Ashi-Kita-cho, limestone rocks by roadside by west side of Kuma river, mossy rocks between both parents, ca. 800 m alt., 1 Nov 1968, Asplenium Sleep & M. Kido AS/605 (holotype, BM (5 fronds); isotype, PE, TI, Z). Planta hybrida tetraploidea, inter parentes A. anogramoideum et A. pekinensem intermedia, sporis abortivis, meiosis valde irregulariter, trivalentibus, bivalentibus univalentibusque.
线裂铁角蕨 xian lie tie jiao jue or 都匀铁角蕨 du yun tie jiao jue
Asplenium bodinieri Christ; A. fuscipes Baker; A. subtoramanum Ching ex S. H. Wu; A. toramanum Makino.
Plants 1235 cm tall. Rhizome erect, short, apex scaly; scales dark brown to black, narrowly triangular to linear-subulate, margin fimbriate to subentire. Fronds caespitose; stipe shiny, purplish black, 615(18) cm, terete, rigid and threadlike, with brown, multicellular uniseriate hairs or subglabrous; lamina triangular, 712 Χ 510 cm, base truncate, bipinnate to tripinnate-pinnatifid, apex acuminate to caudate; pinnae 1015 pairs, subopposite to alternate, lower pinnae stalked, upper almost sessile, stalk dark brown abaxially, basal 1 or 2 pinnae longest and often falcate, narrowly triangular to oblong, 2.57 Χ 1.52 cm, base asymmetrical, on acroscopic side auriculate-truncate, basiscopic side cuneate, up to bipinnate-pinnatifid, apex obtuse; pinnules 68 pairs, anadromous, usually sessile but shortly stalked in more divided fronds, basal acroscopic pinnule largest, ovate, 79 Χ 47 mm, base asymmetrical, acroscopic side truncate, basiscopic cuneate, apex obtuse to truncate; ultimate segments ovate-oblong to linear, apex with 24 short and broadly triangular, obtuse to submucronate or sharp teeth. Costa sulcate adaxially, green, veins often raised adaxially when dry, anadromously branching, (1 or)26 veins per ultimate segment, terminal hydathode obvious, and not reaching margin. Frond firmly herbaceous, green, lamina with multicellular uniseriate hairs or subglabrous; rachis shiny purplish black, becoming green in upper half toward apex, with brown, multicellular uniseriate hairs or subglabrous, sulcate adaxially; costa green except for abaxial side of stalk. Sori (1 or)24 per pinnule or segment, medial to subterminal on acroscopic veinlets, confluent at maturity, oval to linear, 12 mm; indusia grayish green, oval to linear, membranous, repand to entire, opening toward costa or costule, persistent. Spores brown to dark brown, lophate (cristate-alate) perispore. Plants tetraploid: 2n = 144.
On limestone rocks, on man-made constructions[??buildings, walls]; 3001700 m. Fujian, Guangdong, Guizhou, Hunan, Sichuan, Yunnan [Japan, Vietnam].
The type is from China.
Asplenium coenobiale has a variable frond division probably depending on growing conditions with large and more divided plants often growing on rocks in forests. Plants are tetraploid but often produce some aborted spores. Further study will have to show whether they are sexual or agamosporous. In Chinese literature, this name has often been misapplied to the next species (A. pulcherrimum). Due to this confusion, its distribution is not well known.
Chinese pinyin
Davallia pulcherrima Baker, Bull. Misc. Inform. Kew 1895: 53. 1895; Asplenium billetii Christ; A. calcicola Tagawa.
Plants 1025 cm tall. Rhizome erect, apex scaly; scales dark brown to black, narrowly triangular to linear-subulate, margin fimbriate to subentire. Fronds caespitose; stipe shiny, purplish black, 618(25) cm, terete, rigid and threadlike, with brown, multicellular uniseriate hairs or subglabrous; lamina deltoid-triangular, 615 Χ 37(9) cm, base truncate, (3 or)4-pinnate, apex acuminate; pinnae (8)1216 pairs, subopposite to alternate, overlapping, stalk up to ca. 1.3 mm and dark brown abaxially, basal pinnae largest, triangular to ovate, 25 Χ 0.72 cm, base auriculate-truncate, tripinnate, apex obtuse to subacute; pinnules 610 pairs, anadromous, basal acroscopic pinnule largest, triangular-ovate, 512 Χ 58 mm, stalk up to ca. 1 mm, base asymmetrical, acroscopic side truncate, basiscopic side cuneate, apex obtuse, 2-pinnate; ultimate segments 24 pairs, basal acroscopic segment most developed, 24 Χ 24 mm, 2- or 3-partite (fertile) or simple (sterile) and linear-subulate, ultimate fertile segments 0.51.5 mm wide, ultimate sterile segments 0.20.4 mm wide, apex acute. Costa and costules sulcate adaxially, green, veins slightly raised or flat adaxially, anadromous, 1 vein per segment, terminal hydathode not reaching margin. Fronds firmly herbaceous, green, lamina with multicellular uniseriate hairs or subglabrous; rachis shiny purplish black, becoming green in upper half toward apex, sulcate adaxially, with brown, multicellular uniseriate hairs or subglabrous; costa green except for abaxial side of stalk, sulcate adaxially. Sori 1 per fertile, forked and pouch-shaped segment, median on acroscopic veinlet, often spreading and obscuring indusium at maturity, oval, 11.5 mm; indusium white to gray, semi-elliptic, membranous, hyaline, entire, opening toward costules and margin, persistent. Spores brown to dark brown, lophate (cristate-alate) perispore. Plants tetraploid: 2n = 144.
On open rocks or rocks in forests; 7001800 m. Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Sichuan, Taiwan, Yunnan [Malaysia (Sarawak), Vietnam].
The type is from Yunnan.
Plants of Asplenium pulcherrimum are tetraploid but further study will have to show if they are sexual or agamosporous. Due to confusion with the preceding species (Asplenium coenobiale), its distribution is not well known.
Chinese pinyin
Plants 515 cm tall. Rhizome erect, apex scaly; scales dark brown to black, narrowly triangular to subulate, margin fimbriate to subentire. Fronds caespitose; stipe shiny castaneous to purplish black, 0.74 cm, semiterete, threadlike, with brown, multicellular uniseriate hairs or subglabrous; lamina oblong-ovate, 312 Χ 13.5 cm, 3-pinnate, apex acute; pinnae 817 pairs, subopposite to alternate, overlapping, shortly stalked, median pinnae largest, ovate, 0.52 Χ 0.41.5 cm, base truncate, 2-pinnate, apex obtuse; pinnules 24 pairs, anadromous, basal acroscopic pinnule largest, triangular-ovate, stalked, apex obtuse, pinnate; ultimate segments bifid or simple (sterile) and linear-subulate, 12.5 Χ 0.20.5 mm, apex acute; fertile segments wider than sterile. Costa and costules sulcate adaxially, green, veins anadromous, 1 vein per segment, terminal hydathode not reaching margin. Fronds thinly herbaceous, green, lamina with multicellular uniseriate hairs or subglabrous; rachis shiny castaneous to purplish black, becoming green in upper half toward apex, sulcate adaxially, with brown, multicellular uniseriate hairs or subglabrous; costa green, sulcate adaxially. Sori 1 per fertile segment, basal on acroscopic veinlet, often spreading and hidden by indusium at maturity, oval, 0.50.9 mm; indusium semi-elliptic, hyaline, entire to repand, opening toward costules and margin, persistent. Spores brown, with lophate (cristate-alate) perispore.
* In crevices of karst caves; 700800 m. Guangxi.
The type is from Guangxi.
长叶铁角蕨 chang ye tie jue
Asplenium bipinnatum Roxburgh var. prolongatum (Hooker) Bonaparte; A. rutifolium (Bergius) Kunze var. prolongatum (Hooker) Christ.
Plants 2040 cm tall. Rhizome erect, short, apex scaly; scales dark brown to black, narrowly triangular, with narrow pale brown edges, entire or denticulate. Fronds caespitose; stipe green, 818 cm, sulcate adaxially, sparsely covered with small dark brown fimbriate scales when young, later subglabrous; lamina linear-ovate, 1025 Χ 34.5 cm, bipinnate, apex caudate; pinnae 2024 pairs, basal pinnae opposite or subopposite, upper alternate, subsessile, lower pinnae usually not reduced, middle pinnae narrowly elliptic to oblong, 13 Χ 0.81.5 cm, base subsymmetrical, cuneate-truncate, pinnate, apex obtuse; pinnules alternate, anadromous, 25 pairs, linear, 410 Χ 11.5 mm, base adnate to costa, entire, apex obtuse; first acroscopic pinnule 2- or 3-fid. Veins and costa raised adaxially, one vein per pinnule or segment, with terminal hydathode, not reaching margin. Fronds subfleshy but thin when dried, green to yellow-green when dry; rachis green, apex usually prolonged into flagelliform and gemmiferous at apex, flat or shallowly sulcate with raised supravascular ridge on adaxial side, abaxially flat. Sori one per pinnule or segment, median on acroscopic side of subtending vein, linear, 2.55 mm; indusia grayish green, linear, membranous, entire, opening toward costa and margin, persistent.
On tree trunks in forests or on wet rocks; 1002000 m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hainan, Henan, Hubei, SW Hunan, Jiangxi, Sichuan, Taiwan, Xizang, SW Yunnan, Zhejiang [India, Japan, S Korea, Malaysia, Myanmar, Sri Lanka, Vietnam; Pacific islands (Fiji)].
The type is from Japan (Tsushima).
Though Japanese plants of Asplenium prolongatum were reported to be tetraploid with 2n = 144 (Mitui, [??citation]. 1968; Kawakami, [??citation]. 1970); all Chinese plants (Hainan, Guizhou, Guangdong) checked by flow cytometry are hexaploid, and may have to be ascribed to a new taxon. Asplenium Χkenzoi Kurata, the hybrid with A. wrightii, may also occur in China where they often grow together.
岭南铁角蕨 ling nan tie jiao jue
Asplenium tenerum G. Forster var. stenophyllum Bonaparte [stenophilla].
Plants 1535 cm tall. Rhizome erect, short, apex scaly; scales dark brown to black with pale brown narrow edge, triangular to ovate, margin glandular denticulate or fimbriate. Fronds caespitose; stipe 38 cm, semiterete, fleshy, adaxially sulcate but with prominent median supravascular ridge, green or stramineous when dry, base abaxially often dull brown, with dark brown to black triangular scales with filiform apex and smaller hastate-stellate scales; lamina lanceolate, 1325 Χ 25 cm, attenuate to both ends, apex acute-acuminate, bipinnatisect; pinnae 1728 pairs, subopposite or alternate, stalk ca. 1 mm, at base reduced and often triangular, middle pinnae oblong-elliptic, slightly falcate, 1.22.5 Χ 0.61.2 cm, base symmetrical, truncate, pinnatisect, apex obtuse; segments 59 pairs, alternate, anadromous, linear-oblong, 24 Χ 11.5 mm, apex obtuse, at base confluent with costa and forming costal wing, entire, basal acroscopic segment larger and 25-fid. Costa distinct and raised adaxially, obscure abaxially, venation anadromously pinnate, one vein per segment, not reaching margin. Frond usually subfleshy, green to stramineous when dry, with dark brown stellate-hastate scales with filiform tail on abaxial surface; rachis sulcate but with prominent supravascular ridge on adaxial side, green or stramineous when dry, fleshy. Sori one per segment, on acroscopic side of subtending vein, linear, 22.5 mm; indusia whitish to gray-stramineous, oval-linear, membranous, entire, opening toward costa and margin, persistent. Spores with medium to dark brown lophate (costate-cristate) perispore. Plants decaploid: 2n = 360.
On limestone rocks in open forests; 300800 m. Guangdong, Guangxi, S Guizhou, Hainan, SE Yunnan [Vietnam].
The type is from Guangdong.
Asplenium sampsonii is similar to A. thunbergii Kunze [=A. belangeri (Bory) Kunze (1848), not Bory (1833)] from Malaysia and Indonesia, but it is smaller, not gemmiferous on the rachis, the lower pinnae are reduced, and the perispore is different. The distinction between A. sampsonii and A. thunbergii in Chinese floras (e.g., Wu, [??citation]. 1999) is based largely on size differences. Manton ([??citation]. 1954) found that plants from Malaysia are tetraploid, but since specimens from Hainan and Guangxi are decaploid, we are confident that they are different from true A. thunbergii, which probably does not occur in China. Plants of A. tenerum with bipinnatifid fronds can be similar to this species.
膜连铁角蕨 mo lian tie jiao jue
Asplenium caudatum Cavanilles (1802), not G. Forster (1786); A. elongatum Swartz (1806), not Salisbury (1796); A. productum C. Presl; A. tenerum var. terminans Kunze ex Mettenius; Darea tenera (G. Forster) Sprengel.
Plants 3065 cm tall. Rhizome erect, short, apex scaly; scales blackish brown, with pale reddish brown edges, triangular, 35 mm, margin fimbriate to subentire. Fronds caespitose; stipe green or stramineous when dry, 1230 cm, adaxial sulcate with median supravascular ridge, base scaly, or subglabrous; lamina narrowly triangular to linear, 2038(50) Χ 710(14) cm, apex caudate, 1-pinnate to rarely bipinnatifid; pinnae 1525(35) pairs, subopposite to alternate, shortly stalked, lower pinnae not reduced, narrowly triangular, 35 Χ 11.5 cm, simple, base asymmetrical, acroscopic side truncate, basiscopic side cuneate, margin crenate, apex acute or obtuse. Costa stramineous, raised on both sides, venation obvious, pinnate, basal acroscopic vein 24 times forked, other veins simple, not reaching margin. Fronds subleathery, grayish green when dry; rachis stramineous when dry, with hastate-stellate fibrillar scales or subglabrous, sulcate with median supravascular ridge on adaxial side. Sori median on veins linear, ca. 3 mm; indusia whitish to gray-brown, linear, membranous, entire, opening toward costa.
On rocks in dense forests; 4001000 m. Hainan, Taiwan [India, Indonesia, Japan, Korea, Malaysia, Myanmar, Philippines, Sri Lanka, Vietnam; Pacific islands].
The type is from the Pacific islands.
Asplenium tenerum is a variable taxon, probably an aggregate, needing further study. Plants with pinnate-pinnatisect fronds but without reduced basal pinnae most probably belong to this species. Indian plants are tetraploid but the chromosome number of Chinese material is unknown.
骨碎补铁角蕨 gu sui bu jiao jue
Asplenium dareoideum (Mettenius) Makino; A. davallioides Hooker (1857), not Tausch (1839); Humata dareoidea Mettenius.
Plants 2040 cm tall. Rhizome erect, short, apex scaly; scales dark brown, narrowly triangular, margin fimbriate. Fronds caespitose, dimorphic, sterile fronds reduced in size and division; stipe green, base dull purplish or green, 722 cm, base scaly, upward with hairlike scales or subglabrous, sulcate and with supravascular ridge and narrow lateral wings on adaxial side; lamina triangular to broadly triangular-ovate, 1117 Χ 57 cm, apex acuminate or caudate, tripinnate-pinnatifid; pinnae 1012 pairs, alternate, stalked, basal pair largest, narrowly triangular, 37.5 Χ 1.63 cm, base broadly cuneate, subsymmetrical, 2-pinnate, apex acute-acuminate; pinnules 59 pairs, anadromous, acroscopic pinnules larger than basiscopic, ovate-triangular, 12.5 Χ 0.71.3 cm, base cuneate and decurrent on pinnule stalk, pinnate, apex subacute; ultimate segments 4 or 5 pairs, basal acroscopic segment largest and 58 Χ 36 mm, 2- or 3-lobate, ultimate segments oblong, up to 2 mm, furcate or simple, apex subobtuse to mucronate or acute. Costa and veins with raised supravascular ridge on adaxial side, one vein per segment, not reaching margin. Frond subfleshy when living, thin, green when dry; rachis raised on both sides, narrowly winged. Sori one per segment, distal on subtending vein and about as long as subtending segment but not reaching its apex, oval to linear, 24 mm; indusia yellowish brown, semi-elliptic, membranous, margin with reddish glands, often rolling back at maturity, opening toward its costule and margin. Plants tetraploid: 2n = 144.
On rocks in lowland forests; 5001900 m. Fujian, Guangdong, Guizhou, Hainan, Jiangxi, Taiwan, Zhejiang [Japan, Korea].
The type is from Taiwan.
We were unable to trace Metteniuss type specimen of Humata dareoidea in Berlin (B). If it belongs to Asplenium ritoense, then Makinos combination is to be used. Asplenium Χshikokianum Makino, the hybrid between A. ritoense and octoploid A. wrightii, is not uncommon where the parents grow together (e.g., Guizhou, Hunan, Sichuan, and Taiwan).
台南铁角蕨 tai wan tie jiao jue
Asplenium mertensianum Kunze.
Plants 5090 cm tall. Rhizome shortly creeping to erect, scaly. Stipe terete to semiterete, sulcate toward rachis, ca. 40 cm, base dull brown or green, with pale brown, narrowly triangular scales, 810 Χ 1.52 mm with basal margin fimbriate. Lamina ovate-oblong, 4050 Χ 2225 cm, apex acute, 3-pinnate or 4-pinnatifid; pinnae 1015 pairs, lower pinnae opposite or subopposite, close to each other, stalk 0.61 cm, upper pinnae alternate, 13 pairs of basal pinnae not reduced, ovate-triangular, 1217 Χ 37 cm, base broadly cuneate, 2-pinnate, apex acute; pinnules 911 pairs, anadromous, stalked, basal pair longest, narrowly triangular to ovate, 57 Χ 22.5 cm, base cuneate, pinnate, apex subacute; ultimate segments alternate, basal acroscopic segment largest, oblong, 34 Χ 15 mm, bifurcate or simple, apex subacute to obtuse. Veins raised on adaxial surface, one per ultimate segment, not reaching margin. Fronds fleshy, green, with small hastate-stellate scales on abaxial surface; rachis and costa green, sulcate adaxially, with narrow lateral wings. Sori one per pinnule, median on subtending veinlet, oval to linear, 35 mm; indusia grayish, oval to broadly linear, membranous, entire, opening toward costule and margin, persistent. Plants agamosporous, octoploid: 2n = 286.
Taiwan [Japan (Bonin, Ryukyu)].
The type is from S Japan (Bonin).
膜叶铁角蕨属 mo ye tie jiao jue shu
Lin Youxing (林尤兴); Ronald Viane
Herbs, lithophytic, epiphytic, or terrestrial. Rhizome dorsiventral, thin, up to ca. 5 mm in diam., widely creeping, with clathrate scales. Fronds herbaceous, remote; stipe usually shiny and castaneous to dark purplish or black, rarely grayish green, semiterete abaxially, sulcate adaxially; lamina 1-pinnate, rarely simple; rachis sulcate adaxially, basiscopic margin of pinnae often decurrent and forming narrow abaxial wings on rachis; pinnae asymmetrical with basal basiscopic part cut away and becoming dimidiate; basiscopic margin entire, acroscopic margin crenate, undulate, or serrate, sometimes with retuse teeth. Veins free, rarely anastomosing, anadromously branching, becoming simple toward pinna apex, not reaching margin, one to several basal basiscopic veins lacking. Sori solitary, rarely double, linear to subelliptic, indusiate; indusium thinly membranous to papery, free margin entire to erose; stalks of sporangia long uniseriate, annuli of 2028 hardened cells. Spores bilateral, elliptic, perispore elaborate[elaborately ornamented?], exospore smooth; in sexual plants 64 spores per sporangium. Plants sexual or agamosporous. x = (36), 38, 39.
More than 30 species: pantropics; 18 species complexes (eight endemic) in China.
Hymenasplenium was described by Hayata for Asplenium unilaterale Lamarck and based on the peculiar vascular system of its dorsiventral rhizome. It was later reduced to a section of Asplenium by Iwatsuki (Acta Phytotax. Geobot. 27: 44. 1975). Recent molecular phylogenetic analyses show that Hymenasplenium is the most basally diverged genus in Aspleniaceae, not closely related to any of the other members (Murakami, J. Plant Res. 108: 257268. 1995).
Chromosome numbers of Hymenasplenium were first reported, by Manton and Sledge (Philos. Trans., Ser. B, 238: 138, 167. 1954) in H. unilaterale s.l. from Sri Lanka, to be n = 80 and 2n = ca. 158. However, Mitui et al. (Amer. J. Bot. 76: 1691. 1989) showed that the basic chromosome number of Hymenasplenium is x = 39, with one exception being x = 38 in H. subnormale. Apogamous reproduction was discovered in three unrelated Asian species, indicating that such reproductive mode evolved at least three times in the Asian group.
In a cytological study of Hymenasplenium ikenoi (under Asplenium cardiophyllum s.l., Kato et al. (1989[J. Plant Res. 103: 461468. 1990???]) found that its basic number is also x = 39 (n = 78) and concluded that A. cardiophyllum should be included in Hymenasplenium, although this species has cordate simple leaves quite different from those of the other members of Hymenasplenium. The close relation between A. cardiophyllum and Hymenasplenium is supported by molecular studies (Murakami, loc. cit.: page??).
SW China (Yunnan) is the center of diversity of Hymenasplenium. The reports of morphologically intermediate or transitional forms as well as the existence of different cytotypes (ancestral sexual diploids and tetraploids next to agamosporous taxa) in this area, suggest that Hymenasplenium is still in an active state of diversification. Reticulate evolution is not yet reported in this genus but may explain some of the intermediate forms and the difficulty in describing them as clearly separate taxa.
The name (Hymen)asplenium unilaterale has been used erroneously in China for several other taxa, such as H. apogamum, H. hondoense, and H. murakami-hatanakae. True H. unilaterale (Lamarck) Hayata differs in its more strongly dimidiate pinnae, lacking (5 or)7 or 8 basal basiscopic veins, its teeth that are not retuse, and the dark blackish rachis color that distinctly continues onto the costa abaxially. We have not so far found true H. unilaterale occurring in China. All Asian-Himalayan taxa require further study to discover their relationships with Chinese taxa. In particular, Hymenasplenium rivulare (Fraser-Jenkins) Viane & Dong, comb. nov. (Basionym: Asplenium rivulare Fraser-Jenkins, Taiwania 53: 190. 2008, nom. nov. for Asplenium hindusthanense Bir, nom. inval. (Fern Gaz. 14: 309. 1994), proposed as a new name for Asplenium unilaterale var. rivale Beddome) from S India needs attention, as Beddomes (citation?? 1876) drawing (t. 356) shows a venation pattern and marginal retuse teeth with veins ending just below the minute notch, which are characteristic for the E Asian H. retusulume-H. latidens group.
1a. Frond simple .......................................................................................... 1. H. cardiophyllum
1b. Frond pinnate.
2a. Sorus with a double indusium: an inner below sporangia and an outer covering them 18. H. bivalvatum
2b. Sorus with a single indusium, overlying sorus.
3a. Sori terminal on its subtending vein and situated in marginal teeth ............ 2. H. cheilosorum
3b. Sori medial or inframedial, not in marginal teeth.
4a. Marginal teeth entire, rarely (semi)retuse, veins ending just below marginal teeth.
5a. Rachis abaxially dull, grayish green to brown.
6a. Pinnae usually less than 10(15) pairs ........................................... 11. H. subnormale
6b. Pinnae usually more than 15 pairs ..................................................... 12. H. obscurum
5b. Rachis abaxially shiny, castaneous to purplish black.
7a. Middle pinnae with usually more than 3 basal basiscopic veins lacking; dark rachis color abaxially extending onto basal part of costa.
8a. Lamina usually broadest at base, up to 18 cm wide; rachis abaxially blackish purple to black 16. H. excisum
8b. Lamina widest near middle, less than 10 cm wide; rachis abaxially purplish castaneous ........... 17. H. obliquissimum
7b. Middle pinnae with up to 3 basal basiscopic veins lacking, dark rachis color abaxially extending into stipicel[??okay, used thru-out this ms, or change to stalk?? or whatever is best], rarely along costa (in H. hondoense).
9a. Pinnae spreading or deflexed, obtuse ........................................... 15. H. apogamum
9b. Pinnae ascending, subacute-acute.
10a. Stipes more than 5 mm apart; pinnae falcate, sori supramedial-marginal; 1 or 2 basal basiscopic veins missing; 64 spores per sporangium ................... 13. H. murakami-hatanakae
10b. Stipes less than 5 mm apart; pinnae trapeziform to narrowly triangular-falcate, sori medial; 2 or 3 basal basiscopic veins missing; 32 spores per sporangium ........... 14. H. hondoense
4b. Marginal teeth retuse to emarginate due to a (shallow or deep) notch at their apex, most veins ending just below these notches.
11a. Margin shallowly and regularly incised, deepest incisions equal to less than 4 Χ deeper than notch above veins .............................................................................. 3. H. wuliangshanense
11b. Margin more deeply and irregularly incised, deepest incisions usually more than 4 Χ deeper than notch above veins.
12a. Marginal teeth 2.53.5 mm and obscurely retuse ................................ 4. H. retusulum
12b. Marginal teeth up to 2 mm and distinctly retuse.
13a. Stipe and rachis floccose-scaly, pinnae acute ............................... 5. H. furfuraceum
13b. Stipe and rachis subglabrous, pinnae subacute to truncate-obtuse.
14a. Pinnae lunate-falcate, apex subacute to acute, rarely obtuse.
15a. Acroscopic pinna margin bicrenate, rachis abaxially purplish castaneous 9. H. adiantifrons
15b. Acroscopic pinna margin bicrenate-serrate, rachis
abaxially purple
..................................................................................... 10.
H. changputungense
14b. Pinnae trapezoidal, apex usually truncate or obtuse, rarely subacute.
16a. Pinnae spreading, up to 20 pairs; stipe castaneous-purple ........... 8. H. quercicola
16b. Pinnae ascending, more than 20 pairs; stipe dark purplish.
17a. Sori inframedial; pinna margin coarsely crenate-serrate; purple rachis color continued into stipicel abaxially .................................................................. 6. H. szechuanense
17b. Sori medial; pinna margin regularly bicrenate-serrate; purple rachis color not continued into stipicel abaxially ........................................................................... 7. H. latidens
细辛膜叶铁角蕨 xi xin mo ye tie jiao jue
Micropodium cardiophyllum Hance, J. Bot. 21: 268. 1883; Antigramma cardiophylla (Hance) Tardieu; Asplenium cardiophyllum (Hance) Baker; Boniniella cardiophylla (Hance) Tagawa, Phyllitis cardiophylla (Hance) Ching.
Plants up to 30 cm tall. Rhizomes long creeping, with scales and some yellowish brown hairs; scales blackish brown, ovate-triangular, small, margins sparsely toothed, and caducous on older rhizome parts. Fronds well separated, simple, thinly papery, dark to brownish green when dry, subglabrous; stipe shiny, dark brown to black, 1020 cm, terete but with adaxial groove, base with scales and hairs; lamina simple, ovate, 914 Χ 59 cm, base cordate, margin entire or shallowly sinuate, apex acute to acuminate. Midrib (rachis) obvious abaxially, shiny and black to about middle of leaf, lateral veins anadromous, slender, and hardly visible on adaxial side, occasionally connected and forming elongate areolae near margin, vein ends free. Sori linear, usually solitary on acroscopic veins, rarely opposite; indusia persistent, brownish, linear, thinly membranous, entire. Spores elliptic, average exospore length 2124 ΅m, perispore with narrow crests (alae). Plants sexual and diploid.
On rocks or sandy soils in forests near streams. Hainan [Vietnam (Cao Bang)].
The type is from Hainan.
Hymenasplenium cardiophyllum is often classified under Boniniella Hayata (Bot. Mag. (Tokyo) 41: 709. 1927), an endemic genus of E Asia. Recent flow cytometric and cytological studies show that this taxon includes two cytologically different species: tetraploid Hymenasplenium ikenoi (Makino) Viane, comb. nov. (Basionym: Scolopendrium ikenoi Makino, Bot. Mag. (Tokyo) 13: 130. 1899), from Japan (Ryukyu Islands and Bonin Islands), and diploid H. cardiophyllum from China. Apart from their different chromosome number these taxa can also be separated by their average exospore length (2124 ΅m for diploid H. cardiophyllum vs. 2832 ΅m for tetraploid H. ikenoi. However, further studies need to unravel their exact relationships. Plants from Vietnam and Thailand (Boonkerd & Pollawatn, citation??. 2012) require further study as to their ploidy level and true identity, as spore size of the Thai plants suggest a hexaploid situation.
A study of its dorsiventral dictyostele, raphides, chromosome number, and perispore, led Kato et al. ([J. Plant Res. 103: 461468??] 1990) to include H. ikenoi in Hymenasplenium. The chromosome number of 2n = 156 (Kato et al., loc. cit.: page?) indicates that H. ikenoi is a tetraploid with a basic chromosome number of x = 39; this recent count is in agreement with the n = 78 bivalents that can be counted on the photograph published by Kurita (citation??. 1972). A base chromosome number of x = 38 or 39 coincides with that of Hymenasplenium.
Kato et al. (loc. cit.: page?) suggested a closer affinity of H. ikenoi to H. excisum, H. apogamum, and H. obscurum than to H. cheilosorum and the group of H. cataractarum, H. hondoense, and H. obliquissimum. However, recent molecular studies seem to show that H. ikenoi is more closely related to the H. hondoense, H. apogamum, and H. cataractarum clade, and that it originated recently from the ancestor of the above three species (Murakami, J. Plant Res. 108: 257268. 1995).
齿果膜叶铁角蕨 chi guo mo ye tie jiao jue
Asplenium cheilosorum Kunze ex Mettenius, Abh. Senckenberg. Naturf. Ges. 6: 133. 1859; A. heterocarpum Wallich ex Hooker (1859), not Blume (1828).
Plants 2560 cm tall. Rhizome long creeping, 2.54 mm in diam., apex scaly; scales narrowly triangular to triangular, entire or sparsely fimbriate at base. Fronds up to ca. 5 mm apart, dark green when dry, membranous-herbaceous, subglabrous; stipe grayish black to dark purplish, shiny, 1020 cm, base with scales similar to those on rhizome; lamina 1-pinnate, narrowly oblong-triangular, 1535 Χ 35(7) cm, truncate at base, acuminate-caudate at apex; rachis shiny, dark purplish; pinnae 2540 pairs, almost sessile, rectangular to trapeziform or lunate, 1.82.5(3.6) Χ 0.50.9 cm, dimidiate, apex obtuse, base asymmetrical, acroscopic side truncate to cuneate and parallel to rachis, basiscopic side cut away and narrowly cuneate and entire, acroscopic margin deeply crenate to dentate with lobes (ca. 1 mm wide) cut up to 1/42/5 of way to costa, lower pinnae spreading or deflexed, upper pinnae ascending. Veins distinct, forking and free, one vein per marginal lobe and ending below sharp tooth, basiscopic side with 6 or 7 veins lacking. Sori linear, 13 mm, at tip of subtending veins and located in marginal teeth; indusia persistent, yellowish brown to deep brown, semi-elliptic, membranous, entire. Plants sexual diploids (2n = 78), or agamosporous triploids with n = 2n = 117.
In soil or on wet rocks along streams in forests; 5001800 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Taiwan, Xizang, Yunnan, Zhejiang [Bhutan, India, Indonesia, Japan, Malaysia, Myanmar, Nepal, Philippines, Sri Lanka, Thailand, Vietnam].
The type is from Sri Lanka.
As presently circumscribed, Hymenasplenium cheilosorum is an aggregate of a widespread triploid agamosporous and a probably more local (Yunnan) sexual (diploid) taxon. This complex was examined cytologically by Mehra and Bir (Cytologia 25: 1727. 1960) using Himalayan material and later again by Mitui et al. (Amer. J. Bot. 76: 16891697. 1989) using Japanese plants. Mehra and Bir (loc. cit.) listed E Himalayan H. cheilosorum as a triploid agamosporous plant but with the wrong chromosome number of n = 108 (based on x = 36). Mitui et al. (loc. cit.: 16901691) showed that both Himalayan and Japanese H. cheilosorum are apogamous and triploid, but with n = 2n = 117 (based on x = 39) chromosomes. Cheng and Murakami (J. Plant Res. 111: 495500. 1998) found an ancestral sexual diploid taxon in Yunnan, with spore mother cells showing 39 bivalents at metaphase I, but without describing it formally as a separate species. Species of this alliance can be distinguished from the closely related, also sexually reproducing, H. inthanonense N. Murakami & J. Yokoyama from N Thailand, by their more terminal sori and more dimidiate pinnae.
无量山膜叶铁角蕨 wu liang shan mo ye tie jiao jue
Basionym: Asplenium wuliangshanense Ching, Acta Phytotax. Sin. 10: 186. 1965.
Plants 2030 cm tall. Rhizome long creeping, 1.52 mm in diam., apex scaly; scales narrowly triangular to triangular. Fronds remote, up to 10 mm apart, green to brown-green when dry, herbaceous, subglabrous; stipe shiny, dark purplish black, 510 cm, base with scales similar to those on rhizome or subglabrous; lamina 1-pinnate, narrowly oblong-triangular, 1219 Χ 3.55 cm, truncate at base, acuminate to caudate at apex; rachis shiny, purple to purplish black; pinnae 2030 pairs, almost sessile, rectangular to trapeziform or slightly lunate, middle pinnae 1.62.6 Χ 0.50.7 cm, dimidiate, apex obtuse, base asymmetrical, acroscopic side truncate to cuneate and parallel to rachis, basiscopic side cut away and narrowly cuneate and entire, acroscopic margin crenate-serrate with deeply retuse teeth; pinnae spreading, basal pinnae slightly reduced. Veins distinct, forking and free, each vein ending below a marginal notch, basiscopic side with 3 or 4 veins lacking. Sori medial to inframedial, elliptic, 25 mm; indusia persistent, yellowish brown to deep brown, semi-elliptic, membranous, entire, opening toward costa.
* On shaded wet rocks by streams; 19002700 m. W Yunnan.
Hymenasplenium wuliangshanense is most similar to the following taxa (H. furfuraceum to H. changputungense). The upper margin of its pinnae is finely and regularly denticulate, with relatively deep notches (often as deep as the teeth) so that there is no obvious difference between a notch and a normal sinus. H. latidens has slightly coarser teeth and more medial sori. It is also similar to H. adiantifrons, which has a more coarsely cut margin and a more castaneous-purple rachis. Future research will have to establish the true relationships in this group. This and the following six species form a group of related taxa that can best be distinguished by their veins that terminate below a notch in each marginal tooth. A similar taxon, Asplenium ofeliae Soldago, is known from the Philippines.
微凹膜叶铁角蕨 wei ao mo ye tie jiao jue
Basionym: Asplenium retusulum Ching, Acta Phytotax. Sin. 10: 185. 1965.
Plants 2030 cm tall. Rhizome long creeping, 1.53 mm in diam., apex scaly; scales narrowly triangular to triangular. Fronds remote, up to 12 mm apart, green when dry, membranous-herbaceous, subglabrous; stipe shiny, dark purplish black, 412 cm, base with yellowish brown indument, some scales similar to those on rhizome, or subglabrous; lamina 1-pinnate, narrowly oblong-triangular, 1520 Χ 34.5 cm, truncate at base, apex acuminate; rachis shiny, purplish; pinnae 1825 pairs, almost sessile, rectangular to trapeziform, middle pinnae 2.22.6 Χ 0.50.7 cm, dimidiate, apex obtuse, base asymmetrical, acroscopic side truncate to cuneate and parallel to rachis, basiscopic side cut away and narrowly cuneate and entire, acroscopic margin with deep crenate incisions (12 Χ ca. 1 mm) with lobes minutely retuse; pinnae spreading, basal pinnae slightly reduced. Veins distinct, forking and free, one vein per marginal lobe, vein ending below marginal notch, basiscopic side with 3 or 4 veins lacking. Sori inframedial, elliptic, 34 mm; indusia persistent, yellowish brown to brown, semi-elliptic, membranous, entire, opening toward costa. Plants sexual.
* On rocks in mixed forests; ca. 2000 m. SE Yunnan.
Hymenasplenium retusulum is similar to the following taxa from which it differs in its more deeply cut acroscopic margins with relatively long and obtuse teeth, which are only minutely retuse at their tips. Due to confusion with similar taxa, the relationships and the distribution of this species are not well known.
绒毛膜叶铁角蕨 ron mao mo ye tie jiao jue
Basionym: Asplenium furfuraceum Ching, Acta Phytotax. Sin. 10: 190. 1965.
Plants 2030 cm tall. Rhizome long creeping, 1.52 mm in diam., apex scaly; scales narrowly triangular to triangular. Fronds remote, up to 12 mm apart, green when dry, herbaceous, subglabrous; stipe slightly shiny, purplish castaneous, 4.512 cm, with orange-brown woolly-floccose indument of reduced scales, or subglabrous; lamina 1-pinnate, narrowly oblong-triangular, 1219 Χ 3.55 cm, base truncate and slightly reduced, apex acuminate to caudate; rachis shiny and purple to purplish castaneous, with similar indument to that of stipe or subglabrous; pinnae 2025 pairs, almost sessile, color of rachis entering base of stipicel abaxially, trapeziform, middle pinnae 1.62.6 Χ 0.50.9 cm, dimidiate, apex acute, base asymmetrical, acroscopic side truncate and parallel to rachis, basiscopic side cut away and narrowly cuneate and entire, acroscopic margin crenate-serrate with retuse teeth; pinnae spreading to slightly erect, basal pinnae slightly reduced and deflexed. Veins distinct, forking and free, each vein ending below a marginal notch, basiscopic side with 3 or 4 veins lacking. Sori medial, elliptic, 24.5 mm; indusia persistent, brown, semi-elliptic, membranous, entire, opening toward costa.
* On rocks by streams. W Yunnan.
Hymenasplenium furfuraceum differs from similar taxa by its more densely floccose rachis and stipe. Due to confusion with similar species, the relationships and distribution of this species are not well known. Sporangia contain 64 spores, thus the plants are probably sexual.
天全膜叶铁角蕨 tian quan mo ye tie jiao jue
Basionym: Asplenium szechuanense Ching, Acta Phytotax. Sin. 10: 188. 1965.
Plants 2540 cm tall. Rhizome long creeping, 1.52.5 mm in diam., apex scaly; scales dark brown to blackish, narrowly triangular to triangular. Fronds remote, up to 13 mm apart, green to brownish dark green when dry, herbaceous, subglabrous; stipe shiny, dark purplish, 715 cm, base with narrow scales similar to those on rhizome, with brownish indument or subglabrous; lamina 1-pinnate, narrowly oblong-triangular, 1525 Χ 35 cm, base truncate, apex acuminate to caudate; rachis shiny and purple to castaneous-purple, subglabrous; pinnae 2025 pairs, almost sessile, color of rachis entering stipicel and base of costa abaxially, trapeziform to trapeziform-lunate, middle pinnae 1.52.3 Χ 0.71 cm, dimidiate, apex truncate to obtuse, base asymmetrical, acroscopic side truncate and parallel to rachis, basiscopic side cut away and narrowly cuneate and entire, acroscopic margin coarsely crenate-serrate with retuse teeth; pinnae erect, basal pinnae slightly reduced. Veins distinct, forking and free, each vein ending below a marginal notch, basiscopic side with 3 or 4 veins lacking. Sori inframedial, elliptic, 34.5 mm; indusia persistent, brown, semi-elliptic, membranous, entire, opening toward costa.
* On rocks by streams. Sichuan, Yunnan.
Hymenasplenium szechuanense is similar to H. furfuraceum from which it differs in its much less floccose to subglabrous rachis and stipe, its inframedial sori, and its more reddish purple rachis. It differs from H. quercicola in its inframedial sori and more dark purple stipe. Hymenasplenium latidens is also similar but differs by the characters given in the key and by its slightly more numerous pinnae pairs; future research will have to show their relationship. All these taxa also resemble H. obliquissimum but that species has its veins ending in marginal teeth, not below a notch in a marginal tooth as in this group. Due to confusion with similar taxa, the relationships and the distribution of this species are not well known.
阔齿膜叶铁角蕨 kuo chi mo ye tie jiao jue
Basionym: Asplenium latedens Ching, Acta Phytotax. Sin. 10: 187. 1965.
Plants 2540 cm tall. Rhizome long creeping, 1.52.5 mm in diam., apex scaly; scales dark brown, narrowly triangular to triangular. Fronds remote, up to 13 mm apart, green to brownish dark green when dry, herbaceous, subglabrous; stipe shiny, dark purplish, 910(15) cm, base with narrow scales similar to those on rhizome, with brownish indument or subglabrous; lamina 1-pinnate, narrowly oblong-triangular, 1525 Χ 35 cm, base truncate, apex acuminate to caudate; rachis shiny, purple, subglabrous; pinnae up to 30 pairs, almost sessile, color of rachis not entering stipicel abaxially, trapeziform to trapeziform-lunate, middle pinnae 22.6 Χ 0.50.9 cm, dimidiate, apex truncate to obtuse, base asymmetrical, acroscopic side truncate and parallel to rachis, basiscopic side cut away and narrowly cuneate and entire, acroscopic margin shallowly and regularly bicrenate-serrate with retuse teeth; pinnae ascending, basal pinnae slightly reduced. Veins distinct, forking and free, each vein ending below a marginal notch, basiscopic side with 3 or 4 veins lacking. Sori medial, elliptic, 24 mm; indusia persistent, brown, semi-elliptic, membranous, entire, opening toward costa.
* On rocks; 18001900 m. SE Yunnan (Wenshan).
Due to confusion with similar taxa, the relationships and the distribution of Hymenasplenium latidens are not well known. Hymenasplenium szechuanense is very similar but differs by the characters given in the key and above.
镇康膜叶铁角蕨 zhen kang mo ye tie jiao jue
Basionym: Asplenium quercicola Ching, Acta Phytotax. Sin. 10: 188. 1965.
Plants 2035 cm tall. Rhizome long creeping, 1.52.5 mm in diam., apex scaly; scales dark brown to blackish, narrowly triangular to triangular. Fronds remote, up to 12 mm apart, green to brownish dark green when dry, herbaceous, subglabrous; stipe shiny, castaneous-purple, 714 cm, base with scales similar to those on rhizome, or subglabrous; lamina 1-pinnate, narrowly oblong-triangular, 1522 Χ 35 cm, base truncate and slightly reduced, apex acuminate to caudate; rachis shiny, purple to castaneous-purple, pinnae 1520 pairs, almost sessile, color of rachis not entering base of stipicel abaxially, trapeziform, middle pinnae 1.52.5 Χ 0.71 cm, dimidiate, apex truncate to obtuse, base asymmetrical, acroscopic side truncate and parallel to rachis, basiscopic side cut away and narrowly cuneate and entire, acroscopic margin crenate-serrate with retuse teeth, pinnae spreading, basal pinnae slightly reduced and deflexed. Veins distinct, forking and free, each vein ending below a relatively shallow marginal notch, basiscopic side with 3 veins lacking. Sori medial, linear, 34 mm; indusia persistent, brown, semi-elliptic, membranous, entire, opening toward costa.
* Epiphytic on Quercus trees; 24002700 m. Yunnan.
Hymenasplenium quercicola is most similar to H. szechuanense from which it differs in its more castaneous-purple stipe, more spreading pinnae, and more medial sori. Several sporangia of the type have an orange color and appear collapsed or not fully developed, typical features of hybrids in this family. Due to confusion with similar species, the relationships and the distribution of this taxon are not well known.
阿里山膜叶铁角蕨 a li shan mo ye tie jiao jue
Basionym: Asplenium resectum Swartz f. adiantifrons Hayata, Icon. Pl. Formosan. 4: 226. 1914; Asplenium adiantifrons (Hayata) Ching.
Plants 1840(45) cm tall. Rhizome long creeping, 1.52.5 mm in diam., apex scaly; scales narrowly triangular to triangular. Fronds remote, up to 12 mm apart, green to brownish dark green when dry, herbaceous, subglabrous; stipe shiny, castaneous-purple, 520 cm, base with scales similar to those on rhizome or subglabrous; lamina 1-pinnate, narrowly oblong-triangular, 1025 Χ 25 cm, base truncate and slightly reduced, apex acuminate to caudate; rachis shiny, castaneous-purple; pinnae 2030 pairs, almost sessile, color of rachis not entering base of stipicel abaxially, trapeziform to slightly lunate, middle pinnae 1.22.2 Χ 0.50.7 cm, dimidiate, apex subacute to obtuse, base asymmetrical, acroscopic side truncate and parallel to rachis, basiscopic side cut away and narrowly cuneate and entire, acroscopic margin crenate with retuse teeth; pinnae spreading, basal pinnae slightly reduced and deflexed. Veins distinct, forking and free, each vein ending below relatively shallow marginal notch, basiscopic side with 3 or 4 veins lacking. Sori inframedial, linear, 34 mm; indusia persistent, brown, semi-elliptic, membranous, entire, opening toward costa.
* SC Taiwan.
Our description of Hymenasplenium adiantifrons differs from Chings which was solely based on the type and the drawing of Hayata. Although Ching (Acta Phytotax. Sin. 10: 187188. 1965) compared this taxon essentially to H. latidens, it is morphologically also similar to H. changputungense and H. szechuanense. In the literature, this species has sometimes been synonymized with H. filipes [author?] (Philippines) from which it differs, i.a., by its retuse marginal teeth (entire in H. filipes), its shiny stipe and rachis (almost dull in H. filipes), and spreading pinnae (pointing upward in H. filipes).
贡山膜叶铁角蕨 gong shan mo ye tie jiao jue
Basionym: Asplenium changputungense Ching, Acta Phytotax. Sin. 10: 188. 1965.
Plants 2840 cm tall. Rhizome long creeping, 23 mm in diam., apex scaly; scales dark brown, narrowly triangular to triangular. Fronds remote, up to 12 mm apart, green to brownish dark green when dry, herbaceous, subglabrous; stipe shiny, dark purple, 1020 cm, with yellowish hairs or subglabrous; lamina 1-pinnate, narrowly oblong-triangular, 1218 Χ 2.54.5 cm, base truncate and slightly reduced, apex acuminate to caudate; rachis shiny, purple, with yellowish hairs or subglabrous; pinnae 2025 pairs, almost sessile, color of rachis entering stipicel and base of costa abaxially, trapeziform to slightly lunate or upper pinnae subfalcate, middle pinnae 1.32.7 Χ 0.40.7 cm, dimidiate, apex subacute to obtuse, base asymmetrical, acroscopic side truncate and parallel to rachis, basiscopic side cut away and narrowly cuneate and entire, acroscopic margin bicrenate to biserrate with retuse teeth; pinnae spreading to ascending, basal pinnae slightly reduced and deflexed. Veins distinct, forking and free, vein ending below relatively shallow marginal notch, basiscopic side with (3 or)4 veins lacking. Sori inframedial to medial, linear, 2.54 mm; indusia persistent, brown, semi-elliptic, membranous, entire, opening toward costa.
* Mountain slopes; ca. 2000 m. NW Yunnan.
Hymenasplenium changputungense is morphologically closest to H. adiantifrons, H. szechuanense, and H. furfuraceum. However, the relationships and the distribution of these taxa are not well known and need further study.
小膜叶铁角蕨 xiao mo ye tie jiao jue
Asplenium subnormale Copeland in Perkins, Fragm. Fl. Philipp. 183. 1905.
Plants 1520 cm tall. Rhizome long creeping, ca. 3 mm in diam., apex scaly; scales narrowly triangular to triangular. Fronds remote, up to 5 mm apart, brown grayish green when dry, herbaceous, subglabrous; stipe slightly shiny, dark purple-brown to black abaxially, 56 cm, sparsely scaly to subglabrous; lamina 1-pinnate, ovate-triangular, 912 Χ 56 cm, base truncate and slightly reduced, apex acute; rachis grayish green, base slightly shiny and purple, with scales or subglabrous; pinnae up to 10 pairs, distinctly stalked, stipicel broadly falcate-trapeziform, up to 0.3 cm, middle pinnae 2.53 Χ 0.81.3 cm, dimidiate, apex obtuse, base asymmetrical, acroscopic side semicordate to truncate and parallel to rachis, basiscopic side of basal pinnae excavate, in middle pinnae becoming narrowly cuneate and entire, acroscopic margin serrate, teeth not retuse; pinnae spreading to ascending, basal pinnae slightly reduced and deflexed. Veins distinct, forking and free, ending below marginal tooth, basiscopic side with 1(or 2) vein(s) lacking. Sori medial, linear, 26 mm; indusia persistent, pale brown, semi-elliptic to linear, membranous, entire, mainly opening toward costa.
Wet places in valley forests, terrestrial, among weeds by streams, steep forest slopes, mossy rocks in shaded and humid ravines; near sea level to 300 m. S and SE Taiwan, Yunnan [Indonesia, Japan, Malaysia, Philippines].
The type is from the Philippines.
The description of Hymenasplenium subnormale is based on the Yunnan material, which might represent an undescribed species. The type of H. subnormale is a smaller plant in general and differs also essentially from other members of Hymenasplenium in its venation pattern with none of the basiscopic veins lacking (isotypes in MICH!, B!). Morphological variation within this taxon has been pointed out by Holttum (citation?? 1967) and Iwatsuki (Acta Phytotax. Geobot. 27: 51. 1975; citation?? 1995). Mitui et al. (Amer. J. Bot. 76: 1691. 1989) reported a base chromosome number of x = 38 instead of the normal x = 39 for most of the other species in the genus. They also found both diploids (n = 38, 2n = 78) and tetraploids (n = 78) in a collection from Ceram (Indonesia). Cheng and Murakami (J. Plant Res. 111: 495500. 1998) reported a related diploid taxon (A. latipinnum nom. nud.) with n = 39 from Yunnan, different by its more inframedial sori, but without formally publishing it as a new taxon. This taxon is thus clearly an aggregate and plants from outside of the Philippines may represent one or more related species. The whole complex needs a thorough revision.
绿杆膜叶铁角蕨 lό gan mo ye tie jiao jue
Asplenium obscurum Blume, Enum. Pl. Javae 2: 181. 1828; A. erosodentatum Blume; A. obscurum var. angustum Tagawa.
Plants 2040 cm tall. Rhizome long creeping, 34.5 mm in diam., apex scaly; scales dark brown, narrowly triangular to triangular. Fronds remote, up to 10 mm apart, grayish green when dry, papery to herbaceous, subglabrous; stipe dull green to grayish green when dry, not shiny nor purple, 20(25) cm, sparsely scaly to subglabrous; lamina 1-pinnate, narrowly ovate-triangular, 2030 Χ 510 cm, base truncate, apex acuminate to caudate; rachis grayish green, subglabrous; pinnae 2030 pairs, sessile to obscurely stalked, trapeziform-falcate, middle pinnae 3.57 Χ 0.81.3 cm, dimidiate, apex obtuse to subacute, base asymmetrical, acroscopic side truncate and parallel to rachis, basiscopic side of basal pinnae excavate, in middle pinnae narrowly cuneate and entire, acroscopic margin serrate to crenate, teeth not retuse, pinnae spreading to ascending. Veins distinct, forking and free, ending below a marginal tooth, basiscopic side with (3 or)4(or 5) veins lacking. Sori medial, linear, 35 mm; indusia persistent, pale brown, semi-elliptic to linear, membranous, entire, opening toward costa.
Wet places in dense forests, on rocks along streams; 1001600 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Taiwan, Yunnan [India, Indonesia, Myanmar, Nepal, Sri Lanka, Thailand, Vietnam; Africa].
The type is from Java.
Mitui et al. (Amer. J. Bot. 76: 16901691. 1989) reported that this species is a sexual tetraploid with 2n = 156 in Java (Indonesia). Cheng and Murakami (J. Plant Res. 111: 495500. 1998) and Kato et al. (citation?? 1992) reported related sexual diploid and tetraploid taxa from Yunnan (see also under H. bivalvatum); consequently, the taxon as here circumscribed may be a species complex with different cytotypes involved. Contrary to literature citations, Asplenium serriforme Mettenius is not listed here as a synonym because the drawing and the description by Mettenius (citation?? 1859) clearly state that it has retuse (serraturae obtusae emarginato) teeth, and thus represents another taxon probably more related to the H. adiantifrons group.
Reviewer Ralf Knapp notes that the material seen from Taiwan has an inner and outer indusium and might, therefore, match the taxon here described as H. bivalvatum.
单边膜叶铁角蕨 dan bian mo ye tie jiao jue
Asplenium cataractarum Rosenstock (1915), not Blume (1828), H. cataractarum N. Murakami.
Plants 2540 cm tall. Rhizome shortly creeping, 23 mm in diam., apex densely scaly; scales castaneous to dark brown, narrowly triangular; phyllopodia distinct, ca. 2 mm, ca. 10(40) mm distant. Fronds grayish green when dry, herbaceous; lamina 1-pinnate, narrowly triangular to lanceolate, 1020 Χ 35 cm, widest in basal 1/3, gradually narrowing toward apex, glabrous; stipe shiny, purple, 1025 Χ 0.050.15 cm, subglabrous, base sparsely scaly; rachis shiny, purplish abaxially, adaxial side grooved and with 2 grayish green narrow wings; pinnae almost sessile to shortly stalked, purple rachis color extending via stipicel up to 3 mm onto basal part of costa abaxially, 1525 pairs, alternate, usually ascending, falcate to trapeziform, 23.5 Χ 0.61 cm, base asymmetrical, acroscopic side truncate, basiscopic side attenuate-cuneate with ca. 1/2 cut away, acroscopic margin serrate, teeth usually not retuse, pinna apex (sub)acute. Veins forking and terminating in marginal teeth, only 1 or 2 basal basiscopic veins lacking. Sori linear, 23 mm, supramedial to (sub)marginal, indusia brownish, linear, membranous, entire, opening toward costa. Plants diploid, sexual with 2n = 78.
Usually on wet rocks near or in streams in forests. Jiangxi, Taiwan, Yunnan [Japan].
Due to the previous confusion in the H. unilaterale s.l. group, the distribution of Hymenasplenium murakami-hatanakae is not well known. As an ancestral diploid, it may have played an important role in the reticulate evolution of Hymenasplenium in Asia and along the Pacific. However, the elucidation of relationships in the unilaterale group will require much more micromorphological and molecular studies. True H. unilaterale may be largely absent from India, the Himalaya, and the Pacific region, where its nomenclature urgently needs to be brought in line with data published by Cheng and Murakami (citation?? 1998).
Asplenium pubirhizoma Ching & Z. Y. Liu, described from Sichuan, may represent a species of its own although it was considered a synonym of A. unilaterale s.l. by Wu (FRPS 4(2): 38. 1999); it is similar to H. murakami-hatanakae but differs in its gray, hairy scales, and its more medial sori.
东亚膜叶铁角蕨 dong ya mo ye tie jiao jue
Asplenium hondoense N. Murakami & Hatanaka, J. Fac. Sci. Univ. Tokyo, Bot. 14: 191. 1988.
Plants 2540 cm tall. Rhizomes shortly creeping, 23 mm in diam., apex densely scaly; scales castaneous to dark brown, narrowly triangular; phyllopodia distinct, ca. 2 mm tall, ca. 10 mm distant. Fronds grayish green when dry, herbaceous; lamina 1-pinnate, narrowly triangular to lanceolate, 1020 Χ 35 cm, widest in basal 1/3, gradually narrowing toward apex, glabrous; stipe shiny, purple, 1025 Χ 0.050.15 cm, subglabrous, base sparsely scaly; rachis shiny, purplish abaxially, adaxial side grooved and with 2 grayish green narrow wings; pinnae almost sessile to shortly stalked, rachis color extending via stipicel onto base of costa abaxially, 1525 pairs, alternate, usually ascending, trapeziform to falcate, 23.5 Χ 0.61 cm, base asymmetrical, acroscopic side truncate, basiscopic side attenuate-cuneate with 1/31/2 cut away, acroscopic margin serrate, teeth not retuse, pinna apex subacute to obtuse. Veins forking and terminating in marginal teeth, 2 or 3(or 4) basal basiscopic veins lacking. Sori linear, 45 mm, medial; indusia brownish, linear, membranous, entire, opening toward costa. Plants usually agamosporous.
Usually on wet rocks near streams in forests. Fujian, Guangxi, Sichuan [Japan, Korea, Nepal].
The distribution of Hymenasplenium hondoense is not well known due to confusion with other members of the genus. As presently circumscribed, this is a cytologically variable, aggregate species. Usually plants are agamosporous diploids (n = 2n = 72) or triploids (n = 2n = 117), but in China, sexual diploids and tetraploids exist. The elucidation of this aggregate and its relationships to the other SE Asian members of the unilaterale group will require more study.
无配膜叶铁角蕨 wu pei mo ye tie jiao jue
Asplenium apogamum N. Murakami & Hatanaka, J. Fac. Sci. Univ. Tokyo, Bot. 14: 193. 1988.
Plants 2540 cm tall. Rhizomes shortly creeping, 23 mm in diam., apex densely scaly; scales dark brown, narrowly triangular, entire; phyllopodia distinct, ca. 2 mm tall, 23 mm distant from each other. Fronds grayish green when dry, herbaceous; lamina 1-pinnate, narrowly triangular to lanceolate, 1020 Χ 35 cm, widest in basal 1/3, gradually narrowing toward apex, glabrous; stipe shiny, purple, 1025 Χ 0.050.15 cm, subglabrous, base sparsely scaly; rachis shiny and purplish abaxially, adaxial side grooved and with 2 grayish green narrow wings; pinnae almost sessile to shortly stalked, rachis color extending via stipicel onto base of costa abaxially, 1525 pairs, alternate, spreading or deflexed near stipe, quadrangular-trapeziform, 23.5 Χ 0.61 cm, base asymmetrical, acroscopic side truncate and parallel to rachis, subauriculate, basiscopic side attenuate-cuneate, with 1/31/2 cut away, acroscopic margin serrate to sinuate, teeth not retuse, pinna apex obtuse to subacute. Veins forking and terminating in marginal teeth, only 1 or 2 basal basiscopic veins lacking. Sori linear, 34 mm, medial; indusia brownish, linear, membranous, entire, opening toward costa.
Usually terrestrial along streams, broad-leaved forests, shaded areas, moist places; 200500 m (in Taiwan). Taiwan, Yunnan [Japan (Ryukyu Islands), Laos, Thailand, Vietnam].
As circumscribed, Hymenasplenium apogamum is an aggregate species, which, according to Cheng and Murakami (J. Plant Res. 111: 495500. 1998), is both morphologically and cytologically variable. In China (Yunnan), it consists of sexual diploids (n = 39), sexual tetraploids (n = 78), and the more usual and widespread agamosporous triploids (n = 2n = 117). In their 1998 publication (loc. cit.), Cheng and Murakami also recognized another triploid agamosporous taxon (H. laterepens nom. nud) related to H. apogamum, but differing in its more triangular leaves. As the distribution and relationships of the Asian unilaterale group are not well understood, the present treatment only reflects the current state of limited knowledge.
切边膜叶铁角蕨 qie bian mo ye tie jiao jue
Asplenium excisum C. Presl, Epimel. Bot. 74. 1851; A. rahaoense Y. Yabe ex Matsumura & Hayata[IPNI nomen; TROP: nom. nud.]; A. resectum Swartz var. rahaoense Hayata; A. subresectum Copeland; A. unilaterale Lamarck f. majus C. Christensen; A. unilaterale var. majus (C. Christensen) Sledge; A. unilaterale var. rahaoense (Hayata) Hayata; Hymenasplenium rahaoense (Hayata) H. Itτ ex Tuyama.
Plants 4060 cm tall. Rhizome long creeping, 35 mm in diam., apex scaly; scales blackish brown, narrowly triangular to triangular. Fronds remote, up to 8 mm apart, grayish to dark green when dry, herbaceous, subglabrous; stipe shiny and dark purple to black, 1532 cm, sparsely scaly to subglabrous; lamina 1-pinnate, narrowly triangular to triangular, 2040 Χ 1018 cm, base truncate and widest, apex acuminate to caudate; rachis shiny, purplish black, subglabrous; pinnae 1825 pairs, shortly stalked, blackish brown rachis color extending via stipicel onto costa abaxially, trapeziform-oblong to falcate, middle pinnae 58(10) Χ 1.31.8(2) cm, dimidiate, apex acute, base asymmetrical, acroscopic side truncate and often almost parallel to rachis, basiscopic side of basal pinnae excavate, in middle pinnae narrowly cuneate and entire, acroscopic margin serrate, teeth not retuse; pinnae spreading to ascending. Veins distinct, forking and free, ending below a marginal tooth, basiscopic side (3 or)46 veins lacking. Sori medial, linear, 46 mm; indusia persistent, pale brown, semi-elliptic to linear, membranous, entire, opening toward costa.
Shaded and wet places in dense forests, on rocks by streams, on tree trunks; 2001700 m. Guangdong, Guangxi, Guizhou, Hainan, Taiwan, Xizang, Yunnan [India, Malaysia, Myanmar, Philippines, Sri Lanka, Thailand, Vietnam; tropical Africa].
The type is from the Philippines (Luzon).
Hymenasplenium excisum represents a distinct species complex with sexual diploids (2n = 78) and tetraploids (2n = 156) in China (Yunnan) and Indonesia (Mitui et al., Amer. J. Bot. 76: 1691. 1989). Cheng and Murakami (J. Plant Res. 111: 495500. 1998) mention a new endemic species complex, H. costarisorum, from two localities in S Yunnan, and said to be similar to H. excisum in gross morphology but different in smaller size, larger number of pinnae, more strongly reflexed lower pinnae, and sori restricted to the distal part of the pinnae. This undescribed taxon containing two cytotypes: a sexual diploid (n = 36) and a sexual tetraploid (n = 72), has a basic chromosome number of x = 36, which is universal in Aspleniaceae, except Hymenasplenium (with x = 38 and x = 39). Molecular data (Murakami, J. Plant Res. 108: 257268. 1995) showed that this species is closely related to H. obscurum (with a chromosome number of x = 39).
荫湿膜叶铁角蕨 yin shi mo ye tie jiao jue
Asplenium unilaterale var. obliquissimum Hayata, Icon. Pl. Formosan. 4: 230. 1914; A. obliquissimum (Hayata) Sugimoto & Sa. Kurata; A. resectum Swartz var. obliquissimum (Hayata) Hayata; A. unilaterale Lamarck var. udum Atkinson ex C. B. Clarke; Hymenasplenium unilaterale var. obliquissimum (Hayata) Hayata ex Sasaki.
Plants 2030 cm tall. Rhizomes shortly creeping, 1.52.5 mm in diam., apex densely scaly; scales grayish brown, broadly ovate-triangular; phyllopodia distinct, ca. 2 mm tall, up to 10 mm distant from each other. Fronds very thin to translucent, brown-green when dry; lamina 1-pinnate, narrowly triangular to lanceolate, 1524 Χ 2.55 cm, widest in basal 1/3, apex gradually narrowing, subglabrous; stipe shiny, dark castaneous-purple, 1020 cm, subglabrous, base sparsely scaly; rachis shiny, dark castaneous-purple abaxially, adaxial side grooved and with 2 grayish green narrow wings; pinnae strongly dimidiate, shortly stalked, purplish rachis color extending via stipicel onto costa abaxially, 1525 pairs, alternate, ascending, trapeziform to falcate, 1.53.5 Χ 0.40.7 cm, base asymmetrical, acroscopic side truncate, basiscopic side straight with more than 1/2 cut away, acroscopic margin (bi)crenate-sinuate, teeth obtuse and not retuse, apex subacute to obtuse. Veins forking and terminating in marginal teeth, (2 or)3 or 4 basal basiscopic veins lacking. Sori linear, 34 mm, basal on subtending vein and appearing subcostal; indusia brownish, linear, membranous, entire, opening toward costa. Plants sexual or agamosporous.
On rocks near or in streams, in forests; 8002800 m. Guangdong, Guangxi, Guizhou, Hunan, Jiangxi, Sichuan, Taiwan, Yunnan [India, Indonesia, Japan, Nepal, Vietnam].
The type is from Taiwan.
Hymenasplenium obliquissimum is an aggregate of species with different chromosome number and life strategy; three cytotypes, based on x = 39, are known. A diploid sexual (2n = 78), a tetraploid sexual (2n = 156), and a triploid agamosporous type (2n = 117) which is probably a hybrid between the diploid and the tetraploid (Murakami, citation?? 1989).
In the literature, this species has sometimes been put into synonymy with H. filipes (Philippines) but considering cytological and morphological differences, we prefer to keep these taxa distinct. Hymenasplenium obliquissimum differs from H. filipes by its thinner lamina, its thicker rhizome (up to 0.15 cm in H. filipes), its shinier stipe and rachis (almost dull in H. filipes), the rachis color extending onto the costa, its larger number of pinnae (up to 15 in H. filipes), its larger number of reduced basal basiscopic veins (1 or 2 in H. filipes), and its sorus position along the subtending vein (median in H. filipes).
双盖膜叶铁角蕨 shuang gai mo ye tie jiao jue
Basionym: Asplenium unilaterale Lamarck var. bivalvatum Nayar & Geevarghese, Fern Fl. Malabar, 292. 1993.
Plants 3050 cm tall. Rhizome long creeping, 35 mm in diam., apex scaly, scales narrowly triangular to triangular. Fronds remote, 510 mm apart, grayish to dark green when dry, herbaceous, subglabrous; stipe shiny and dark purple to black abaxially but green adaxially, 1020(25) cm, base scaly to subglabrous; lamina 1-pinnate, narrowly triangular to ovate-oblong, 2030 Χ 810 cm, base truncate, apex acuminate to caudate; rachis shiny, purplish black, subglabrous; pinnae 2030 pairs, shortly stalked, trapeziform-oblong to falcate, middle pinnae 37 Χ 0.81.5 cm, dimidiate, apex subacute, base asymmetrical, acroscopic side truncate and often almost parallel to rachis, basiscopic side of basal pinnae excavate, in middle pinnae narrowly cuneate and entire, acroscopic margin serrate, teeth not retuse. Veins distinct, forking and free, ending below marginal tooth. Sori medial, linear, 1.53 mm; indusia persistent, double, consisting of an outer and an inner part, pale brown, semi-elliptic to linear, membranous, entire, opening toward costa. Plants sexual, diploid with 2n = 78.
Shaded forest floor near water courses; ca. 500 m. Taiwan, Yunnan [India, Thailand, Vietnam].
The type is from India.
Cheng and Murakami (J. Plant Res. 111: 495500. 1998) used the name bilabiatum for this taxon, but did not publish a valid name. According to these authors, Hymenasplenium bivalvatum is a distinct species in SW China, and seems to be related to H. excisum. Because Kato et al. (citation?? 1992) reported 2n = 156 (4x) with a sexual reproductive mode for the same species from the same locality, this probably is an aggregate of a diploid and a tetraploid species. The special soral structure with an inner and an outer indusium has never been recorded in this or related fern families. Murakami (J. Plant Res. 108: 257268. 1995) reported it from N Thailand to N Vietnam and Taiwan.