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

1. Arachniodes Blume, Enum. Pl. Javae. 2: 241. 1828.

复叶耳蕨属 fu ye er jue shu

Authors: He Hai, Sugong Wu, Xiang Jianying & David S. Barrington

Byrsopteris C. V. Morton; Leptorumohra (H. Itô) H. Itô; Lithostegia Ching; Phanerophlebiopsis Ching; Rumohra sect. Leptorumohra H. Itô.

Plants terrestrial, evergreen, medium-sized (or large). Rhizome long creeping to short and ascending, or rarely erect, densely scaly; scales filiform, subulate, lanceolate, or ovate, often attenuate toward apex, margin entire or with toothlike (rarely fimbriate) projections. Fronds few, remote or approximate; stipe equal in length to lamina or longer, base as densely scaly as rhizome, adaxially sulcate; lamina deltoid, ovate, pentagonal, or rarely lanceolate, usually 2-4(or 5)-pinnate, but simply pinnate in a few species, herbaceous, papery to leathery, glabrous or with a few narrow scales, rarely hairy; dissection anadromous throughout; rachis adaxially sulcate, grooves continuous with those on axes of higher order, scaly to glabrescent, or rarely with pale gray unicellular acicular hairs, scales entire (or with irregular projections on margin), sometimes with enlarged and denticulate base; pinnae a few to numerous pairs, usually shortly stalked, basal ones often with long basal basiscopic pinnules; upper divisions gradually reduced and ending in acuminate or caudate apices, or with a distinct terminal pinna; basiscopic pinnules often slightly longer than acroscopic ones; ultimate segments sessile, usually oblong to rhomboid, or rarely linear, often inequilateral, with an acroscopic auricle, margin sharply dentate to aristate; venation usually also anadromous, veins free, pinnate or forked, not quite reaching margin of ultimate segment, often ending in a hydathode. Sori orbicular, terminal or dorsal on acroscopic branches of veinlets; indusia orbicular-reniform, attached at a deep sinus, persistent or fugacious; sporangia numerous, long stalked, with an annulus of 13-16 thickened cells. Spores ellipsoid, often with broad rugate perispore. x = 41.

About 60 species: tropical and subtropical regions worldwide, but mainly in E and SE Asia; 40 species (18 endemic) in China.

The following taxon is excluded from the present treatment, pending further research: Arachniodes heyuanensis Ching (Bull. Bot. Res., Harbin 6(3): 9. 1986), described from Guangdong.

1 Rhizome slender and long creeping; lamina herbaceous, rather thin and usually soft; axes and main veins adaxially with pale gray unicellular acicular hairs   (2)
+ Rhizome usually stout, creeping to ascending, or erect; lamina papery to leathery, or rarely herbaceous especially when young; axes and main veins adaxially glabrescent or with scales, but not unicellular hairs   (4)
2 (1) Indusium entire; main veins abaxially with broadly ovate and bullate-based scales; stipe and primary rachis with copious scales.   1 A. miqueliana
+ Indusium ciliate or entire; main veins abaxially with narrowly lanceolate, or hairlike and flat-based scales; upper part of stipe and primary rachis glabrescent   (3)
3 (2) Lamina pentagonal, 4(or 5)-pinnate, upper parts conspicuously abruptly narrowed toward apex and acuminate, brownish green when dried.   2 A. quadripinnata
+ Lamina deltoid-pentagonal, 3-pinnate-pinnatifid, upper parts long acuminate, yellowish green when dried.   3 A. sinomiqueliana
4 (1) Lamina 4- or 5-pinnate, ultimate segments very narrowly lanceolate, with a single vein; sori solitary in each segment; indusia very firm, dark and entire, covering sorus when young and splitting into irregular segments when mature.   4 A. superba
+ Lamina usually 2-4(or 5)-pinnate, rarely simply pinnate, ultimate segments asymmetrically oblong or rhomboid, with more than one vein; sori often more than one in each segment; indusia membranous, papery or firm, persistent or fugacious, margin entire or erose   (5)
5 (4) Lamina oblong-lanceolate, simply pinnate; pinnae lanceolate; free ultimate segments (pinnae) nearly symmetrically cuneate at base   (6)
+ Lamina deltoid, ovate, or pentagonal, 2-4(or 5)-pinnate; pinnae usually deltoid or ovate-lanceolate; free ultimate segments usually asymmetrical at base with acroscopic auricles   (8)
6 (5) Stipe with dark brown scales at base; pinnae rachis abaxially with sparse hairlike scales; sori mostly dorsal at or below middle of veinlets.   5 A. neopodophylla
+ Stipe with reddish brown scales at base; pinnae rachis abaxially glabrescent; sori terminal or subterminal on veinlets   (7)
7 (6) Lamina acuminate at apex or with a distinct terminal pinna; pinnae with aristate-tipped cartilaginous coarse teeth on margin.   6 A. blinii
+ Lamina with a distinct terminal pinna; pinnae with non-aristate-tipped crenulate teeth on margin.   7 A. tsiangiana
8 (5) Sori dorsal on veinlets; ultimate segments with margin dentate or acutely serrate, never with aristate tips; texture thickly leathery.   8 A. cavaleriei
+ Sori terminal on veinlets (or subdorsal but mixed with terminal ones on same lamina); ultimate segments with margin acutely dentate or pinnatilobate, teeth with or without aristate tips; texture papery, subleathery, or herbaceous   (9)
9 (8) Rhizome and base of stipe with shiny castaneous, ovate-lanceolate, and stiffly thickly papery scales; stipe upward and rachis throughout glabrescent and glossy   (10)
+ Rhizome and base of stipe with blackish, grayish, or reddish brown, ovate-lanceolate, lanceolate, linear-lanceolate, or filiform, and membranous or thinly papery scales; stipe upward and rachis throughout ?scaly and often not glossy   (11)
10 (9) Lamina caudate toward apex, and with a distinct terminal pinna; lateral pinnae also caudate at apices.   9 A. amoena
+ Lamina abruptly narrowed toward apex and caudate-acuminate, without distinct terminal pinna; lateral pinnae also caudate-acuminate at apices.   10 A. tonkinensis
11 (9) Rhizome ascending, with tufted, reddish brown, long, tortuous, filiform scales extending to stipe base   (12)
+ Rhizome long to shortly creeping or ascending, often bearing grayish or blackish brown, linear-lanceolate, ovate-lanceolate, or subulate scales extending to stipe base   (16)
12 (11) Lamina 2-pinnate-pinnatifid, free ultimate segments 5-8 ?ca. 1.5 cm   (13)
+ Lamina 3-5-pinnate, free ultimate segments at most to 2 ?1 cm   (14)
13 (12) Stipe base with only long hairlike scales; lamina leathery, lustrous and brownish when dried.   11 A. grossa
+ Stipe base with broad, denticulate-margined scales and with hairlike scales adaxially; lamina papery, dull dark green when dried.   12 A. fengii
14 (12) Fronds large, usually reaching ca. 1.7 m; lamina 5-pinnate; stipe with both subulate scales and broadly lanceolate, denticulate-margined thicker scales along both sides of adaxial groove at lower part above base.   15 A. gigantea
+ Fronds medium-sized, at most to 1.5 m; lamina 3- or 4-pinnate; stipe with scales similar to those on rhizome or bearing slightly wider scales along both sides of adaxial groove near base   (15)
15 (14) Lamina papery, 3- or 4-pinnate; ultimate segments obviously separate from each other, apex and lobes mucronate, dull brownish when dried; indusia usually denticulate on margin.   13 A. globisora
+ Lamina thinly leathery, 4-pinnate; ultimate segments contiguous to imbricate, apex and lobes with non-aristate sharp teeth, shiny green when dried; indusia often entire on margin.   14 A. spectabilis
16 (11) Rhizome lignified, together with base of stipe bearing linear-lanceolate, lanceolate, or subulate, and thicker and firmer scales   (17)
+ Rhizome succulent, especially at growing apex, bearing broadly lanceolate or ovate-lanceolate and more delicate scales extending to stipe base   (28)
17 (16) Fronds dimorphic; fertile lamina strongly contracted with much shorter and narrower pinnae and pinnules.   16 A. hainanensis
+ Fronds monomorphic, or with minor differences between fertile and sterile laminae (or pinnae on same lamina)   (18)
18 (17) Lamina abruptly acuminate to caudate-acuminate toward apex, most often with a conform terminal pinna resembling lateral pinnae   (19)
+ Lamina gradually acuminate to strongly acuminate, gradually reduced without a conform terminal pinna   (21)
19 (18) Lamina 2- or 3-pinnate, with an elongated caudate-acuminate apex, lateral pinnae often linear-lanceolate (only lowest pinnae deltoid-lanceolate).   19 A. longipinna
+ Lamina (2 or)3- or 4-pinnate, with a distinct terminal pinna resembling adjacent lateral pinnae, lower 2-4 pairs of pinnae often deltoid-lanceolate with long basal basiscopic pinnules   (20)
20 (19) Lamina relatively smaller and less dissected when mature, (2 or)3-pinnate, lower pinnae proximally with 1 or 2 pairs of elongated and 1-pinnate pinnules.   17 A. simplicior
+ Lamina relatively larger and more dissected, 4-pinnate at base, lower pinnae proximally with 2 or 3 pairs of elongated and 1- or 2-pinnate pinnules.   18 A. ziyunshanensis
21 (18) Lamina 2- or 3-pinnate; stipe bearing brown or dark brown, linear-lanceolate to subulate, denticulate-based, appressed scales above base and rachis throughout   (22)
+ Lamina (3 or)4(or 5)-pinnate; stipe bearing brown or coal-black, linear-lanceolate (or wider but often entire-margined) spreading scales above base and rachis throughout, or stipe glabrescent   (25)
22 (21) Lamina caudate-acuminate at apex; lower pinnae with basally elongated pinnules   (23)
+ Lamina acuminate to shortly acuminate at apex; pinnae (except basal) often without basally elongated pinnules   (24)
23 (22) Rhizome long creeping; lamina shiny brownish green when dried; sori medial on veinlets.   20 A. aristata
+ Rhizome shortly creeping; lamina dull greenish when dried; sori medial or more basal on veinlets.   22 A. jinpingensis
24 (22) Lamina acuminate to attenuate at apex, occasionally abruptly narrowed; lateral pinnae often with basal pinnules longest.   21 A. chinensis
+ Lamina acuminate at apex; lateral pinnae with basal pinnules as long as or shorter than adjacent upper ones.   23 A. japonica
25 (21) Lamina subleathery, stiff or firm when dried   (26)
+ Lamina thinly herbaceous, soft and tenuous when dried   (27)
26 (25) Lamina thickly papery to subleathery when mature; ultimate segments broader and with aristate or mucronate teeth.   24 A. speciosa
+ Lamina papery or thinly papery when mature; ultimate segments often narrower and with only sharply tipped (but non-aristate) teeth.   25 A. coniifolia
27 (25) Stipe above base and rachis throughout bearing sparse, dull brown, relatively thinner scales, or glabrescent.   26 A. festina
+ Stipe above base and rachis throughout bearing a number of coal-black and relatively thicker scales.   27 A. nigrospinosa
28 (16) Lamina abruptly caudate at apex and forming a conform linear terminal pinna   (29)
+ Lamina toward apex acuminate, attenuate, or abruptly narrowed but acute, or forming a broadly acuminate terminal pinna   (31)
29 (28) Sori marginal or submarginal; indusia margin ciliate at least when young.   28 A. amabilis
+ Sori medial; indusia margin entire   (30)
30 (29) Pinnae strongly ascending (forming an angle of less than 30?with rachis), lower 3 or 4 pairs similar in shape and size; ultimate segment margin serrate; indusia with hairs on dorsal surface.   30 A. similis
+ Pinnae obliquely spreading (forming an angle of more than 60?with rachis), pinnae gradually reduced toward lamina apex (and sometimes lowest pair of pinnae with long basal basiscopic pinnules); proximal ultimate segments on acroscopic or both margins often shallowly pinnate; indusia glabrous.   29 A. hekiana
31 (28) Lamina 2-pinnate; ultimate segments often broader than 1 cm   (32)
+ Lamina 2-pinnate-pinnatifid to usually 3- or 4(or 5)-pinnate; ultimate segments often narrower than 1 cm   (35)
32 (31) Lamina gradually acuminate toward apex   (33)
+ Lamina abruptly narrowed toward apex, with a distinct but acuminate terminal pinna   (34)
33 (32) Lamina as long as or shorter than stipe, deltoid-oblong, with a terminal pinna resembling or slightly wider than its lateral ones; ultimate segments not decurrent.   31 A. assamica
+ Lamina longer than stipe, often oblong, its 1-pinnate apex much longer and wider than lateral pinnae; ultimate segments decurrent.   32 A. yoshinagae
34 (32) Stipe ochraceous; lamina papery, brownish green when dried; lateral pinnae broader than 5 cm.   33 A. pseudoassamica
+ Stipe stramineous; lamina thinly papery, yellowish green when dried; lateral pinnae narrower than 4 cm.   34 A. wulingshanensis
35 (31) Lamina abruptly narrowed toward apex and long acuminate or acute; lowest pinnae linear-lanceolate to lanceolate or basal 1(or 2) pairs of pinnules obviously elongated   (36)
+ Lamina acuminate or slightly attenuate at apex; lowest pinnae deltoid or ovate and often much broader, with basal basiscopic pinnules slightly longer   (37)
36 (35) Lamina 2-pinnate-pinnatifid; lateral pinnae linear-lanceolate or lanceolate; ultimate segments often deeply pinnatifid.   35 A. hunanensis
+ Lamina 3-pinnate; lower 3(or 4) pinnae deltoid with elongated basal pinnules; ultimate segments (except proximal ones) sharply serrate.   36 A. sinorhomboidea
37 (35) Lamina thickly papery or subleathery, most often 3-pinnate; ultimate segments usually broader than 5 mm   (38)
+ Lamina herbaceous or papery, most often 4-pinnate; ultimate segments usually narrower than 5 mm   (39)
38 (37) Pinnules remote; lamina papery and often flexible, shiny yellowish green when dried; veins and costules with brown, appressed articulate hairs abaxially.   37 A. nipponica
+ Pinnules contiguous; lamina subleathery and often firm, pale greenish when dried; veins and costules glabrescent abaxially.   38 A. ailaoshanensis
39 (37) Ultimate segments with aristate-tipped teeth; indusia ciliate or erose; lamina firmly herbaceous or papery, often dull pale green when dried.   39 A. simulans
+ Ultimate segments with blunt-tipped teeth; indusia entire; lamina thinly herbaceous, often shiny green when dried.   40 A. henryi

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