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19. Lepisorus (J. smith) Ching, Bull. Fan Mem. Inst. Biol. 4: 47. 1933.

瓦韦属 wa wei shu

Authors: Qi Xinping, Zhang Xianchun, Youxing Lin, Michael G. Gilbert & Peter H. Hovenkamp

Drynaria sect. Lepisorus J. Smith, Bot. Mag. 72. Comp. 13. 1846; Belvisia Mirbel; Drymotaenium Makino; Hymenolepis Kaulfuss (1824), not Cassini (1817); Macroplethus C. Presl; Platygyria Ching & S. K. Wu.

Plants epiphytic or epilithic, rarely terrestrial. Rhizomes creeping, ± terete or slightly flattened, densely scaly when young, sometimes naked when old, sometimes with white waxlike covering; scales blackish brown, opaque or clathrately transparent, ovate, orbicular, or broadly lanceolate to subulate-lanceolate, entire to deeply serrate. Fronds simple, remote or closely spaced, monomorphic, less often ± dimorphic; stipe usually short, base sparsely scaly, upper part smooth, mostly straw-colored, less often dark brown; lamina mostly lanceolate, less often narrowly lanceolate to linear or auriculate to pedately lobed, margin entire or undulate, often revolute when dried, mostly leathery or papery, less often herbaceous when dried, both surfaces glabrous or abaxially sparsely scaly. Midrib obvious, lateral veins often obscure, veinlets reticulate, areoles with simple or forked included veinlets, sometimes with hydathodes at ends. Sori large, orbicular or elliptic, sometimes confluent into linear coenosori, in 1 row on each side of midrib, superficial or sometimes deeply impressed, covered with paraphyses when young; paraphyses peltate, entire or denticulate, less often stellate or scalelike, often brown at center, pale at margin; lumina large, transparent. Sporangia usually leptosporangiate: long stalked, subpyriform, annulus longitudinal, consisting of 14 conspicuously thickened cells; less often sporangia platygyroid: subspherical, annulus of much wider thin-walled cells. Spores ellipsoid, without perispore, surface mostly rugose or undulate, less often smooth, tuberculate, or foveolate. 2n = 39, 46, 50, 52, 70, 74, 94, 95, 100, 148, 150.

About 80 species; mainly in E Asia, a few in Africa, one species in Hawaii; 49 species (23 endemic) in China.

The type species is Lepisorus nudus (Hooker) Ching (Pleopeltis nuda Hooker, described from Nepal), not, as sometimes indicated, L. thunbergianus (Kaulfuss) Ching, which was not included in J. Smith’s original treatment of Drynaria sect. Lepisorus.

Lepisorus is complex both cytologically and morphologically with many taxa still inadequately known. Because of this, we have treated species conservatively. Though relatively recently described, the genus has become well established, especially as molecular studies have shown that the similarity to the genus Pleopeltis Humboldt & Bonpland ex Willdenow is probably the result of morphological convergence and that the two genera belong to phylogenetically distinct lines. These same studies (e.g., Li Wang et al., Molec. Phylogen. Evol. 54: 211-225. 2010) demonstrated that two older, but much smaller, genera, Belvisia and Drymotaenium, nest within Lepisorus. Lepisorus has been formally proposed for conservation against these competing earlier names.


1 Lamina clearly differentiated into a much wider basal vegetative portion and a much narrower apical fertile portion almost completely covered by a pair of linear sori   (2)
+ Lamina mostly with little or no differentiation between vegetative and fertile portions, sometimes sori restricted to apical portion but this only slightly narrower and sori not covering surface   (4)
       
2 (1) Lamina narrowly lanceolate, upper parts gradually attenuate into narrow fertile part, herbaceous when dry.   1 L. mucronatus
+ Lamina ovate-lanceolate or elliptic, upper parts abruptly contracted into narrow fertile part, papery or leathery when dry   (3)
       
3 (2) Stipe very short to almost absent; lamina leathery with undulate margins when dried, fertile tip 2-3 mm wide; sori covering entire abaxial surface of fertile tip when mature.   2 L. henryi
+ Stipe 2-5 cm; lamina papery with straight margins when dried, fertile tip 5-10 mm wide; sori separated from midrib by a narrow sterile zone.   3 L. annamensis
       
4 (1) Sori linear or merged into linear coenosori when mature   (5)
+ Sori orbicular or elliptic, separate   (8)
       
5 (4) Sori a pair of continuous linear coenosori each within a longitudinal groove alongside midrib; lamina linear.   5 L. miyoshianus
+ Sori superficial, not inserted within any grooves, often discontinuous toward base of lamina; lamina lanceolate   (6)
       
6 (5) Rhizome scales 1-1.3 ?0.3-0.45 mm; sori linear, tip of fertile lamina abruptly narrowed.   4 L. sinensis
+ Rhizome scales 2-4 ?0.7-1 mm; sori initially separate, confluent into interrupted coenosori when mature, tip of fertile lamina more gradually narrowed   (7)
       
7 (6) Lamina 0.4-0.8 cm wide; paraphyses with margins colorless, entire or erose.   22 L. subconfluens
+ Lamina 1-2.5 cm wide; paraphyses with margins brown and spiny.   24 L. confluens
       
8 (4) Rhizome long creeping, wirelike, to 1 mm in diam.; lamina 0.3-1.5 cm in width   (9)
+ Rhizome shortly to long creeping, not wirelike, (1-)1.5-4 mm in diam.; lamina lanceolate to linear lanceolate, margin entire, auriculate or laminae hastate, width various   (10)
       
9 (8) Paraphyses to 0.2 mm in diam., very thick, with almost opaque lumina.   6 L. ussuriensis
+ Paraphyses 0.3-0.5 mm in diam., very thin, with transparent square lumina.   7 L. pseudoussuriensis
       
10 (8) Rhizome scales opaque in center, never spreading   (11)
+ Rhizome scales translucent or transparent in center, spreading or adpressed   (28)
       
11 (10) Rhizome shortly creeping, leaves caespitose.   8 L. tosaensis
+ Rhizome long creeping, leaves remote   (12)
       
12 (11) Lamina linear-lanceolate, 2-5 mm wide, margin strongly revolute, protruding where covering sori so frond like a string of beads (moniliform)   (13)
+ Lamina linear-lanceolate to lanceolate, more than 5 mm in width, rarely less than 5 mm, margin not or only slightly revolute and lamina never moniliform   (14)
       
13 (12) Lamina to 15 cm; paraphyses with lumina dense and opaque.   9 L. lewisii
+ Lamina more than 20 cm; paraphyses with lumina square and transparent   10 L. monilisorus
       
14 (12) Paraphyses stellate, lumina dense; rhizome scales without paler margins   (15)
+ Paraphyses orbicular, lumina dense or square; rhizome scales with paler margins   (17)
       
15 (14) Laminae loriform, 40-70 cm, widest at middle.   23 L. medogensis
+ Laminae lanceolate, less than 30 cm, widest 1/3-1/2 of length from base   (16)
       
16 (15) Sori confluent, lamina apex constricted; rhizome scales ?transparent.   24 L. confluens
+ Sori separate, lamina apex not constricted; rhizome scales opaque.   25 L. sordidus
       
17 (14) Stipe base dark; scales thick; lamina sometimes moniliform due to protruding sori   (18)
+ Stipe base straw-colored or chestnut-brown; scales brown to dark brown, thin; lamina not moniliform   (19)
       
18 (17) Lamina linear-lanceolate, apex not constricted.   11 L. heterolepis
+ Lamina lanceolate, apex constricted, sori restricted to constricted part   12 L. luchunensis
       
19 (17) Rhizome scales glossy; lumina of paraphyses square.   13 L. lineariformis
+ Rhizome scales dull and not iridescent; lumina of paraphyses small and polygonal   (20)
       
20 (19) Stipes mostly chestnut-brown; paraphyses 0.15-0.30 mm in diam..   14 L. obscurevenulosus
+ Stipes straw-colored; paraphyses more than 0.3 mm in diam   (21)
       
21 (20) Lamina broadly lanceolate, abaxial surface distinctly scaly; rhizome thick, scales dark brown   (22)
+ Lamina lanceolate, leaf indument often absent; rhizome much slender, scales brown   (23)
       
22 (21) Lamina widest at middle, abaxial surface densely scaly; sori costular, close to each other.   15 L. oligolepidus
+ Lamina widest at lower 1/3, abaxial surface more sparsely scaly; sori medial and remote.   16 L. suboligolepidus
       
23 (21) Rhizome scales ovate-lanceolate, with only a narrow opaque band in center, otherwise transparent   (24)
+ Rhizome scales lanceolate, with a broad opaque band in center part and narrow transparent margins   (25)
       
24 (23) Scales yellow-brown, ?spreading, only with a narrow opaque band at center   17 L. elegans
+ Scales brown, imbricate, with ?opaque band.   18 L. contortus
       
25 (23) Rhizome scales chestnut to black in center, margins serrate; distal sori often confluent.   22 L. subconfluens
+ Rhizome scales brown in center, margins entire or denticulate; sori not confluent   (26)
       
26 (25) Rhizome scale apex not filiform; lamina widest below middle.   19 L. thunbergianus
+ Rhizome scale apex long and filiform; lamina widest at middle   (27)
       
27 (26) Lamina linear, 0.3-0.5 cm wide.   20 L. angustus
+ Lamina lanceolate, 0.5-1.8 cm wide.   21 L. tibeticus
       
28 (10) Rhizome scales with lumina dense and translucent, margin entire, rarely denticulate; paraphyses orbicular   (29)
+ Rhizome scales lanceolate, transparent, clathrate, margin denticulate; paraphyses stellate or lanceolate   (40)
       
29 (28) Rhizome scales lanceolate, persistent, with distinctly paler margins; plants summer green   (30)
+ Rhizome scales ovate or lanceolate, caducous, or if scales persistent then lamina to 4 cm wide, scales not distinctly bicolored; plants evergreen   (32)
       
30 (29) Rhizome scales with orbicular base and acute apex, closely appressed to rhizome, 1-2 mm, not overlapping, rhizome surface exposed, white farinose.   35 L. bicolor
+ Rhizome scales gradually acuminate, overlapping, 2.5-4 mm, rhizome surface hidden   (31)
       
31 (30) Rhizome scales lanceolate, distinctly bicolored, dark brown center contrasting with white margins.   36 L. morrisonensis
+ Rhizome scales broadly lanceolate, margin only slightly paler than center.   37 L. scolopendrium
       
32 (29) Rhizome scales ovate to acuminately ovate, margin entire, lumina isodiametric   (33)
+ Rhizome scales lanceolate or ovate-lanceolate, margin entire or denticulate, lumina rectangular, rarely isodiametric   (36)
       
33 (32) Sori submarginal; scales ovate, thin, 2-3 mm.   26 L. macrosphaerus
+ Sori subcostular or medial; scales ovate to acuminately ovate, thick, less than 2 mm   (34)
       
34 (33) Sori small, close, medial or subcostular, generally extending to below middle of lamina; lamina margins undulate when dry.   27 L. marginatus
+ Sori large, distant, medial, generally not extending below middle of lamina; lamina margins not undulate when dry   (35)
       
35 (34) Lamina 1.5-4 cm wide; rhizome scales 1.2-2 ?1-1.3 mm; paraphyses brown.   28 L. asterolepis
+ Lamina 1-2 cm wide; rhizome scales ca. 1 ?1 mm; paraphyses pale brown.   29 L. kawakamii
       
36 (32) Rhizome scales ovate-lanceolate, margin denticulate; leaves widest at base.   30 L. megasorus
+ Rhizome scales lanceolate, margin entire; leaves widest at or below middle   (37)
       
37 (36) Lamina more than 4 cm wide; rhizome scales pale brown.   31 L. kuchenensis
+ Lamina 1-3 cm wide; rhizome scales brown to dark brown   (38)
       
38 (37) Rhizome scale margin yellow-brown, center dark brown, central lumina small and square.   32 L. subsessilis
+ Rhizome scale margin pale brown, center brown, central lumina large and rectangular   (39)
       
39 (38) Sori nearer costa; leaves less than 30 cm; rhizome 1.5-2 mm in diam..   33 L. nudus
+ Sori nearer margin; leaves up to 65 cm; rhizome 2-4 mm in diam..   34 L. affinis
       
40 (28) Paraphyses stellate, small in diam., lumina small, often opaque; lamina leathery, evergreen, venation hidden, young leaves always red or with red costa   (41)
+ Paraphyses lanceolate, rhomboidal or deeply stellate, margin spiny, lumina uniformly large and transparent; lamina membranous, summer green, venation distinct, lamina never red or partly red   (45)
       
41 (40) Lamina linear, less than 5 mm wide, margin strongly revolute, protruding around mature sori so frond moniliform; paraphyses opaque.   38 L. eilophyllus
+ Lamina linear, or lanceolate, (2-)10-20 mm wide, margin straight, not revolute; paraphyses transparent or opaque   (42)
       
42 (41) Lamina loriform; sori marginal.   39 L. loriformis
+ Lamina lanceolate; sori medial   (43)
       
43 (42) Lamina broadly lanceolate, 2-5 cm wide.   40 L. sublinearis
+ Lamina lanceolate, less than 2 cm wide   (44)
       
44 (43) Rhizome erect; leaves clustered.   41 L. cespitosus
+ Rhizome creeping; leaves remote.   42 L. pseudonudus
       
45 (40) Sporangia with broad, unthickened platygyroid annulus, ?indehiscent   (46)
+ Sporangia with normal, narrow, thickened annulus cells   (47)
       
46 (45) Lamina lanceolate or oblong-lanceolate, never enlarged at base.   48 L. clathratus
+ Lamina hastate, pedatifid, or auriculate at base, sometimes only some fronds enlarged at base.   49 L. waltonii
       
47 (45) Rhizome scales lanceolate, 3.5-7 ?0.7-1.2 mm at base, 2.5-6 ?as long as wide; areoles oblong or irregularly elongated, longer than wide   (48)
+ Rhizome scales broadly lanceolate, 1.8-2.5 ?0.8-1.5 mm, less than 2.5 ?as long as wide; areoles polygonal, ?as long as wide   (50)
       
48 (47) Rhizome with both hairs and scales, ca. 4 mm in diam.; lamina thick, 1.9-2 cm wide   43 L. tricholepis
+ Rhizome with few or no hairs, 1.5-3 mm in diam.; lamina 0.8-1.5 cm wide   (49)
       
49 (48) Lamina membranous or thinly herbaceous, narrowly lanceolate; rhizome scales black, iridescent, 3.5-7 ?0.7-1 mm.   48 L. clathratus
+ Lamina thickly papery, lanceolate; rhizome scales brown, brittle, 3-3.5 ?0.9-1.2 mm.   44 L. crassipes
       
50 (47) Lamina 5-14 ?0.4-1.2 cm; rhizome scales < 2 ?1 mm, paraphysis margins with long thick curved spines.   45 L. albertii
+ Lamina > 14 ?1-2 cm; rhizome scales 2-2.5 ?1.1-1.5 mm, paraphysis margins not like above   (51)
       
51 (50) Lamina papery or membranous, both surfaces brown, pale brown or pale green when dried, stipe 2-10 cm; paraphyses 0.4-0.5 mm, 2-3 ?as long as wide   46 L. thaipaiensis
+ Lamina thickly papery to thinly leathery, abaxially gray-green, adaxially yellow- or dark green, or both surfaces light green when dried, stipe 0.4-5 cm; paraphyses 0.8-1 mm, ?as long as wide.   47 L. likiangensis

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