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5. Diplazium Swartz, J. Bot. (Schrader). 1800(2): 61. 1801.

双盖蕨属 shuang gai jue shu

Authors: He Zhaorong & Masahiro Kato

Allantodia R. Brown; Anisogonium C. Presl; Callipteris Bory; Monomelangium Hayata.

Plants terrestrial, evergreen, medium-sized to large. Rhizome creeping to ascending or erect, sometimes subarborescent, brown or nearly black, dictyostelic, ± scaly; scales subulate, lanceolate, or broadly lanceolate, sometimes ovate and adpressed or peltate, margin entire, sometimes sparsely toothed and often narrow, black, brown, yellow-brown, or dark brown to black; fronds caespitose, distant or approximate. Stipe brown or black and scaly at base, scales similar to those on rhizome, and upward gradually glabrescent or with sparse scales, sometimes base and upward prickly, grooved adaxially, not hairy. Lamina mostly broadly ovate, oblong, or deltoid, sometimes broadly lanceolate, usually pinnate to tripinnate-pinnatifid under acuminate and pinnatifid apex, mostly bipinnate or bipinnate-pinnatilobate to bipinnate-pinnatipartite. Pinnae lanceolate, broadly lanceolate, or oblong-lanceolate, base mostly or nearly symmetrical, lower pinnae not reduced or slightly reduced; ultimate pinnules mostly lanceolate or broadly lanceolate, sometimes ovate or oblong, base stalked or sessile, truncate, rounded, or subcordate, margin entire or serrate, apex acute or acuminate to long acuminate, sometimes rounded or truncate; veins free, rarely anastomosing with a row of areoles along costa and costule; veinlets simple or forked, ascending, lower veinlets often curved. Lamina herbaceous or papery, rarely leathery, usually glabrous, rachis, costae, and costules abaxially sometimes with sparse subulate or lanceolate scales, rarely prickly; rachis, costae, and costules adaxially grooved between knife-edgelike ridges, ridges interrupted at insertion of higher-order frond axes, so grooves continuous, both surfaces of costa often with minute glandular hairs. Sori linear, oblong, or ovate, mostly single or less often double on basal acroscopic veinlets, costular, medial, supramedial, or from base to end of veinlets; indusia membranous or thickly membranous, grayish white or brown, vaulted or ovoid when mature, and irregularly rupturing on back, or compressed, linear, opening acroscopically, persistent or caducous. Spores bilateral, monolete, mostly reniform or bean-shaped, sometimes hemispherical, rarely spherical; perispore generally prominent, ornamentation mostly rugate, glabrous or aculeate, granular, clathrate, reticulate, rarely ridged. n = 41.

About 300-400 species: mainly distributed in the tropics and subtropics worldwide; a few species distributed in warm temperate and temperate zones or subalpine belts; 86 species (29 endemic) in China.

Most of the species in China are distributed south of the Chang Jiang and in SW regions on mountains; they occur in evergreen broad-leaved forests or beside shaded streamlets, and a few extend to 3000 m.

The following species, accepted in Knapp (Ferns Fern Allies Taiwan, 152, 451, 487. 2011), has been excluded from the present treatment: Diplazium proliferum (Lamarck) Thouars (Esquisse Fl. Tristan d’Acugna, 35. 1808; Asplenium proliferum Lamarck, Encycl. 2: 307. 1786; Callipteris prolifera (Lamarck) Bory). Further study is needed to establish its presence in Taiwan.

The following taxon, described from Guizhou, is excluded from the present treatment, pending further research: Diplazium platyphyllum Christ (Bull. Acad. Int. Géogr. Bot. 17: 148-149. 1907).

1 Veins anastomosing   (2)
+ Veins free   (3)
2 (1) Rachis glabrous, not spiny.   1 D. esculentum
+ Rachis spiny.   2 D. paradoxum
3 (1) Stipe, rachis, and costae densely pilose   (4)
+ Stipe, rachis, and costae not densely pilose   (5)
4 (3) Fronds up to 65 cm, basal pinnae not strongly reduced; veins hairy.   3 D. pullingeri
+ Fronds up to 25 cm, basal pinnae reduced; veins subglabrous.   4 D. dinghushanicum
5 (3) Lamina imparipinnate with terminal pinna similar to lateral pinnae   (6)
+ Lamina gradually shallowly pinnatifid to apex   (14)
6 (5) Sori usually single, occasionally double, distant   (7)
+ Sori mostly double, attached back to back, approximate   (8)
7 (6) Lamina subleathery; margin of pinna usually serrate only above middle, subentire or shallowly repand near base.   5 D. crassiusculum
+ Lamina herbaceous; pinna margin serrate from base to apex, sometimes shallowly pinnatilobate.   6 D. pinfaense
8 (6) Pinnae up to 7 cm wide; veinlets 3(or 4) per veinlet group; scales ovate-lanceolate, 2-3 mm wide.   7 D. hainanense
+ Pinnae 2-5.5 cm wide; veinlets (3 or)4 per veinlet group or more; scales linear-lanceolate or lanceolate, up to 1 mm wide   (9)
9 (8) Pinnae serrate, crenate, or repand from base to apex   (10)
+ Pinnae entire except slightly serrate in apical part   (11)
10 (9) Pinnae subleathery, base rounded, margin repand or crenate (crenations ca. 1.5 mm).   8 D. maonense
+ Pinnae papery, base cuneate or rounded-cuneate, margin minutely serrate or repand.   9 D. serratifolium
11 (9) Pinnae ovate-lanceolate, ca. 2 cm wide.   10 D. basahense
+ Pinnae ovate-lanceolate or elliptic-lanceolate, 3-5.5 cm wide   (12)
12 (11) Scales membranous, bicolored (middle brown, margin black); lateral pinnae up to 12 pairs.   11 D. splendens
+ Scales thickly membranous, unicolored; lateral pinnae up to 6 pairs   (13)
13 (12) Rhizome erect; scales linear, black.   12 D. stenolepis
+ Rhizome slender, creeping to oblique; scales lanceolate, brown or black-brown.   13 D. donianum
14 (5) Sori ovate-orbicular or cylindrical-oblong, costular; indusia vaulted, ovate-orbicular or shortly sausage-shaped, irregularly ruptured on back   (15)
+ Sori shortly oblong, narrowly elliptic, shortly or long linear; indusia mostly not vaulted, opening acroscopically   (24)
15 (14) Scales on rhizome apex and stipe base loose, lanceolate or linear-lanceolate   (16)
+ Scales on stipe ovate or ovate-lanceolate, sometimes lanceolate, peltate or subpeltate (with cordate base)   (17)
16 (15) Plants tall; rhizome ascending to erect, robust, more than 5 cm in diam.; stipe above 6 mm in diam.; pinnules usually with broad base, deltoid-lanceolate; veinlets on lobes mostly forked; perispore with prominent spores, with dense granular ornamentation.   14 D. bellum
+ Plants medium-sized; rhizome creeping, stipe less than 3 mm in diam.; pinnules linear-lanceolate, base same width as middle; veinlets on lobes mostly simple; spores without prominent perispore with dense granular ornamentation.   15 D. ovatum
17 (15) Stipe and base of rachis densely scaly when young, scales at apex of variously long stalks, long stalks adpressed (stipe rough)   (18)
+ Scales on stipe sparse, often adpressed, rarely at apices of stiff conical shortly aculeate protuberances, stipe smooth or prickly after scales fallen   (19)
18 (17) Rhizome erect, fronds caespitose; stipe with estipitate scales, also with adpressed uniseriate-celled, small, nodose hairs.   16 D. dulongjiangense
+ Rhizome creeping, fronds distant; stipe lacking uniseriate-celled nodose hairs.   17 D. kawakamii
19 (17) Rhizome thick, ascending to erect; fronds caespitose   (20)
+ Rhizome creeping; fronds sparse   (21)
20 (19) Plants medium-sized; lamina less than 1.5 m; costae glabrous.   18 D. medogense
+ Plants large; lamina more than 3 m; costae often prickly and rough.   19 D. sikkimense
21 (19) Lamina 3-pinnate, ultimate pinnules pinnatilobate.   20 D. muricatum
+ Lamina 2-pinnate, pinnules pinnatilobate   (22)
22 (21) Pinnule lobe entire or subentire.   21 D. doederleinii
+ Pinnule lobe prominently serrate or shallowly pinnatilobate   (23)
23 (22) Pinnule oblong-lanceolate, middle wider, or lanceolate, base nearly symmetrical; middle and lower pinnule lobes usually shallowly lobed, apex shallowly crenate.   22 D. spectabile
+ Pinnule usually deltoid-lanceolate, base wider; pinnule lobes slightly subfalcate, oblique, apex subtruncate, margin serrate, not lobed.   23 D. subspectabile
24 (14) Sori usually broadly oblong, occasionally elliptic or shortly cylindrical; indusia prominently vaulted, easily ruptured   (25)
+ Sori and indusia shortly or long linear; indusia not vaulted, opening acroscopically, often reflexed under sporangia at maturity   (30)
25 (24) Lamina papery, 2-pinnate; pinnules usually shallowly pinnatilobate to pinnatifid, or only serrate; scales always toothed at margin   (26)
+ Lamina herbaceous, 3-pinnate or 2-pinnate with pinnules pinnatipartite to pinnatisect; lobes of secondary pinnule prominently serrate or pinnate; scales entire   (27)
26 (25) Sori supramedial or submarginal, indusia markedly swollen, elliptic or shortly cylindrical.   24 D. conterminum
+ Sori medial or inframedial, indusia slightly swollen, oblong when mature.   25 D. virescens
27 (25) Lamina 2-pinnate; pinnules pinnatipartite; pinnule lobes regularly dentate at margin; sori 2-3 ?ca. 1 mm, densely contiguous when mature; perispore with slightly rugate, granular ornamentation.   26 D. multicaudatum
+ Lamina 3-pinnate or nearly 3-pinnate (pinnules pinnatipartite to pinnatisect, lobes connected with narrow wings); sori mostly ca. 1(-1.5) mm; perispore with aculeate ornamentation   (28)
28 (27) Rhizome creeping; lamina herbaceous or thickly herbaceous, often tall, ca. 2 m, base 3-pinnate; pinnule lobe or secondary pinnule oblong or linear-lanceolate, apex mostly acute to shortly acute, less often rounded.   27 D. prolixum
+ Rhizome creeping or ascending to erect; lamina herbaceous or thinly herbaceous, usually less than 1 m, sometimes up to 1.5 m, base 3-pinnate, sometimes 2-pinnate; secondary pinnule ovate or oblong, apex rounded or acute   (29)
29 (28) Rhizome ascending to erect; lamina herbaceous, secondary pinnule of lower pinna mostly acuminate at apex, serrate at margin.   28 D. pseudosetigerum
+ Rhizome creeping to ascending; lamina thinly herbaceous, secondary pinnule rounded at apex, often pinnatifid to pinnatipartite.   29 D. alatum
30 (24) Lamina 1-pinnate, usually lanceolate, broadly lanceolate, or ovate-lanceolate; pinnae entire or pinnatilobate to pinnatipartite   (31)
+ Lamina of well-grown plants 2-pinnate or base nearly 3-pinnate   (48)
31 (30) Plants small or medium-sized; lamina oblong, broadly lanceolate, or broadly deltoid-lanceolate, rarely ovate-deltoid or subdeltoid; pinna mostly sickle-shaped, lanceolate at base, asymmetrical, acroscopic base prominently auriculate, margin usually serrate or shallowly lobed   (32)
+ Plants medium-sized; lamina oblong; pinnae oblong-lanceolate, base symmetrical or nearly symmetrical, exauriculate protuberances acroscopic, mostly pinnatilobate to pinnatisect at margin   (37)
32 (31) Rhizome slender, creeping; fronds distant   (33)
+ Rhizome short, ascending to erect; fronds caespitose   (35)
33 (32) Stipe base with dense, brown, translucent membranous, entire lanceolate scales; upper part to middle of rachis with many black, lanceolate or narrowly lanceolate, entire thick scales; pinnae shallowly lobed (lobes shallowly serrate at margin), rarely slightly deltoid processes; stalks of pinnae except basal pinnae narrowly winged.   30 D. nigrosquamosum
+ Stipe base with sparse brown, lanceolate, entire, thickly membranous scales   (34)
34 (33) Lamina herbaceous or thinly herbaceous; stipe sparsely scaly; rachis occasionally with black-brown, lanceolate scales; pinnae with shallow deltoid lobes on both sides, lobes shallowly serrate; pinna stalk narrowly winged.   31 D. okudairai
+ Lamina thickly papery or subleathery; upper part of stipe subglabrous; pinnae double- or single-toothed; pinna stalk wingless.   32 D. wichurae
35 (32) Stipe not more than 1 mm in diam.; middle pinnae less than 3 cm, base (including basal auricle) less than 1 cm wide; pinnae mostly contiguous or imbricate, auricle usually overlapping rachis.   33 D. heterocarpum
+ Stipe ca. 2 mm in diam.; middle pinnae (3-)5-8(-11) cm, base (1-)1.5-2(-4) cm; pinnae usually approximate but imbricate, auricle not overlapping rachis   (36)
36 (35) Lamina ovate-deltoid or subdeltoid; pinnae pinnatifid to pinnatisect or base nearly pinnate (pinnatisect with 1-3 pairs of ovate lobes, pinna stalk short, narrowly winged).   34 D. dushanense
+ Lamina broadly oblong-lanceolate; pinnae pinnatilobate or obtusely toothed at margin.   35 D. lobulosum
37 (31) Rhizome long creeping; fronds distant   (38)
+ Rhizome ascending to erect, occasionally creeping; fronds caespitose, rarely approximate and sparse   (40)
38 (37) Lamina acuminate at apex; pinnae usually pinnatilobate to pinnatipartite, sometimes repand or serrate, scales glabrous on surface.   36 D. mettenianum
+ Lamina abruptly narrowed, then acuminate to apex; pinnae dentate at margin, at most shallowly pinnatilobate   (39)
39 (38) Pinnae 10-13 pairs, base rounded; scales linear-lanceolate, superficially shortly hairy.   37 D. hirtisquama
+ Pinnae 6-8 pairs, base mostly shallowly cordate; scales lanceolate, not hairy   38 D. yaoshanense
40 (37) Lamina herbaceous; stipe and rachis densely scaly or persistently scaly; pinnae sessile   (41)
+ Lamina papery or thickly papery; stipe except base and rachis nearly lacking persistent scales; pinnae stalked   (42)
41 (40) Pinnae deeply pinnatilobate or pectinate; pinna lobes regularly crenate at margin.   39 D. hirsutipes
+ Pinnae shallowly pinnatilobate or incised or broadly toothed; lobes entire or subentire.   40 D. hirtipes
42 (40) Lamina nearly 1-imparipinnate, terminal pinna subsimilar to and larger than lateral pinnae, base broadened, pinnatipartite with 1 or many ligulate oblong lobes   (43)
+ Lamina abruptly or gradually narrowed, acuminate at apex   (44)
43 (42) Base of pinna rounded or cuneate, regularly shallowly crenate at margin.   41 D. changjiangense
+ Base of pinna cuneate, bidentate at margin.   42 D. wangii
44 (42) Lamina acuminate at apex; pinnae pinnatipartite to pinnatisect, lobes densely crenate at margin.   43 D. incomptum
+ Lamina abruptly narrowed to acuminate apex; pinnae pinnatilobate to pinnatifid or only serrate; pinna lobes entire or slightly shallowly serrate   (45)
45 (44) Pinnae shallowly pinnatilobate or only incised with broadly serrate margin; pinna stalk short, up to 5 mm in basal pinnae   (46)
+ Lower pinnae pinnatifid to pinnatipartite; stalk of basal pinna more than 1 cm   (47)
46 (45) Pinnae broadly oblong-lanceolate, more than 25 ?5 cm, base mostly shallowly cordate; perispore few rugate.   44 D. megaphyllum
+ Pinnae sickle-shaped, lanceolate, less than 15 ?3 cm, base crenate or nearly so; perispore prominent and with uniform reticulate ornamentation.   45 D. pinnatifidopinnatum
47 (45) Base of pinna often broad, asymmetrical, broadly crenate, acroscopic and basiscopic lobes prominently alternate; perispore with thick anastomosing ornamentation.   46 D. jinpingense
+ Base of pinna slightly narrowed or as wide as middle, symmetrical or nearly symmetrical, cuneate or broadly cuneate, lobes opposite or subopposite; perispore broadly hyaline, few rugate.   47 D. siamense
48 (30) Sori and indusia thin, long linear; pinnae, pinnules, and pinnule lobes mostly short and broad, apical pinnae near lamina apex often not pinnatilobate; pinnules and pinnule lobes entire or shallowly crenate; fertile frond 1- or 2-pinnate at base   (49)
+ Sori and indusia shortly linear or linear-oblong; pinnules and pinnae near lamina apex pinnatipartite or pinnatilobate, at least dentate on all margins   (51)
49 (48) Veins of pinna and pinnule anastomosing with costal deltoid and subcostal rectangular areoles.   48 D. stenochlamys
+ Veins not anastomosing, rarely anastomosing with 1 row of costal areoles   (50)
50 (49) Rhizome creeping or ascending; lower pinnae broadly lanceolate; pinnules and pinnule lobes incised or crenate; veinlets usually simple, occasionally basal veinlets united to form areoles.   49 D. matthewii
+ Rhizome short, erect; lower pinnae oblong or narrowly lanceolate; pinnules and pinnule lobes entire; veinlets mostly forked.   50 D. metcalfii
51 (48) Lamina ovate, usually 2-pinnate; pinnules and pinnule lobes mostly rounded or acute-ovate or narrowly ovate at apex, sometimes pinnules of lower pinnae ovate or broadly deltoid-lanceolate, rarely linear-lanceolate; if lamina 1-pinnate then lobes oblong or ovate-oblong   (52)
+ Pinnules often lanceolate, acuminate or long acuminate at apex; plants robust and tall   (56)
52 (51) Fertile fronds 1- or 2-pinnate at base; scales entire.   51 D. baishanzuense
+ Lamina 2-pinnate   (53)
53 (52) Rhizome usually ascending to erect, rarely also creeping; pinnules pinnatipartite to pinnatifid; scales toothed at margin   (54)
+ Rhizome creeping; pinnule pinnatipartite to pinnatisect   (55)
54 (53) Base of pinnule prominently asymmetrical, auriculate on acroscopic side, cuneate and slightly decurrent to rachis on basiscopic side; stipe base with sparse broadly lanceolate scales, upper part of stipe and rachis without persistent scales.   52 D. kansuense
+ Base of pinnule symmetrical or nearly symmetrical, not auriculate, not decurrent to rachis; stipe and rachis usually with many persistent lanceolate or broadly lanceolate, black-brown scales.   53 D. squamigerum
55 (53) Pinnules of lower pinnae linear-lanceolate, less than 1.5 cm wide, sessile or subsessile; scales broadly lanceolate, sparsely thinly toothed at margin.   54 D. sibiricum
+ Pinnules of lower pinnae ovate or deltoid-lanceolate, 2.5-3.5 cm wide, prominently stalked, up to ca. 5 mm; scales linear-lanceolate, entire.   55 D. leptophyllum
56 (51) Lamina papery, thickly papery, or subleathery, shiny; pinnules usually pinnatilobate to pinnatifid, or margin shallowly serrate or entire, rarely pinnatifid   (57)
+ Lamina mostly herbaceous, not shiny; pinnules mostly pinnatifid to pinnatipartite, pinnule lobes mostly contiguous or pectinate   (66)
57 (56) Pinnules and subapical pinnae entire.   56 D. glingense
+ Pinnules pinnatilobate, at least shallowly incised or serrate at margin   (58)
58 (57) Scales entire; plants medium-sized   (59)
+ Scales toothed at margin; plants large or medium-sized   (60)
59 (58) Lamina 1-pinnate with pinnae pinnatifid to pinnatipartite, or nearly 2-pinnate, or in lower half 2-pinnate with basal 1 or 2 pairs of pinnules in basal pinnae pinnatipartite to pinnatisect, 1 pinnule or 1 pair in middle pinnae adnate or sessile; stipe brown or brown-green.   57 D. petelotii
+ Lower half of lamina 2-pinnate; pinnules adnate to cuneate base; stipe gray-stramineous.   58 D. subdilatatum
60 (58) Scales prominently bicolored with black margin   (61)
+ Scales concolored, margin not black   (63)
61 (60) Stipe and rachis quadrangular-cylindrical, square in cross section; perispore with thick anastomosing ornamentation.   59 D. quadrangulatum
+ Stipe and rachis terete; perispore with non-anastomosing ornamentation   (62)
62 (61) Pinnules lanceolate or ovate-lanceolate, base often asymmetrical, shallowly cordate or broadly cuneate, basiscopic base broader, sometimes subtruncate.   60 D. dilatatum
+ Pinnules linear-lanceolate or narrowly elliptic-lanceolate, base rounded or rounded-cuneate, symmetrical or nearly symmetrical.   61 D. uraiense
63 (60) Plants large; rhizome ascending, up to 5 mm in diam.; stipe and lower half of rachis densely scaly; scales yellow-brown, linear-lanceolate or lanceolate; lamina 2-pinnate, pinnules shortly stalked; sori shortly linear, costular, 1/3-1/2 as long as veinlets.   62 D. forrestii
+ Plants medium-sized; rhizome creeping, occasionally ascending; scales brown or black-brown; middle and upper parts of stipe glabrous; lamina at most 2-pinnate, pinnules sessile or subsessile; sori linear, not costular, more than 3/4 length of veinlets   (64)
64 (63) Pinnae falcate, lanceolate, asymmetrical with basiscopic pinnules longer, free pinnules 1 or 2(or 3) pairs, ligulate-oblong or falcately lanceolate, apex obtuse or acute, base truncate or adnate; veinlets usually 2-forked; sori sparse, ca. 3/4 as long as veinlets.   63 D. griffithii
+ Pinnae broadly lanceolate or lanceolate, mostly not falcate, symmetrical or nearly so; free pinnules 3-5(-8) pairs   (65)
65 (64) Pinnules sickle-shaped lanceolate, pinnatilobate or prominently dentate at margin, veinlets mostly bifurcate; sori broad, shortly linear, dense on pinnular lamina; lamina stiffly papery when dry.   64 D. petrii
+ Pinnules lanceolate, margin shallowly incised or repand, or entire; veinlets mostly simple; sori sparse, thin; lamina thinly papery when dry.   65 D. tibeticum
66 (56) Scales on rhizome apex and stipe base adpressed, or stipe not scaly   (67)
+ Scales on rhizome apex and stipe base loose   (69)
67 (66) Pinnules pinnatilobate to pinnatifid, symmetrical or nearly symmetrical; sori as long as veinlets; stipe and lower part of rachis often with fleshy green aculeate or verrucose processes.   66 D. simile
+ Pinnules pinnatipartite to costa, asymmetrical, basal basiscopic lobes often large and pinnate; sori reaching to 2/3 of veinlet length; stipe and rachis glabrous   (68)
68 (67) Evergreen; rhizome often ascending to erect, up to 40 cm tall, ca. 10 cm in diam., also creeping; lamina thickly herbaceous; pinnules sessile or slightly shortly stalked; perispore prominent, hyaline, with granular ornamentation.   67 D. laxifrons
+ Summer green; rhizome creeping; lamina thinly herbaceous; pinnules mostly long stalked, up to ca. 4 mm; perispore surface with dense aculeate ornamentation.   68 D. taquetii
69 (66) Stipe and rachis or at least rachis with prominent prickly protuberances   (70)
+ Stipe and rachis not spiny, often smooth, sometimes with sparsely verrucose protuberances or green fleshy aculeate protuberances   (71)
70 (69) Stipe glabrous, rachis with thin prickly processes; base of stipe ca. 7 mm in diam., sparsely scaly; scales ovate-lanceolate, rusty membranous; pinnule lobes separate, connected to narrow wings, margin shallowly pinnatilobate or prominently dentate.   69 D. amamianum
+ Stipe and rachis prickly; base of stipe 1.5-2 cm in diam., densely scaly; scales linear or linear-lanceolate, light brown, bicolored, with narrow black margin, up to 2 cm, membranous.   70 D. asperum
71 (69) Scales entire   (72)
+ Scales toothed at margin   (76)
72 (71) Rhizome erect, with stiff, black-brown roots; stipe brown; pinnule lobes broadly oblong or rectangular.   71 D. viridescens
+ Rhizome creeping, roots not stiff   (73)
73 (72) Scales broadly lanceolate, cells of central part of scale wider than those of marginal part; spores globose.   72 D. jinfoshanicola
+ Scales lanceolate or linear-lanceolate, cells of central part of scale as narrow a s those of marginal part; spores reniform or elliptic   (74)
74 (73) Basal part of lamina nearly 3-pinnate with pinnules pinnatisect to costules; scales lanceolate, membranous; sori slender, shortly linear; perispore with aculeate processes.   73 D. chinense
+ Lamina 2-pinnate with pinnules pinnatilobate or pinnatipartite; scales thickly membranous; sori thick, shortly linear; perispore without aculeate processes   (75)
75 (74) Lamina thickly herbaceous; scales lanceolate; pinnules shallowly pinnatilobate or pinnatipartite 1/2 to base; apex of pinnule lobe falcate, truncate or rounded-truncate; perispore with many verrucose projections.   74 D. hachijoense
+ Lamina herbaceous; scales linear-lanceolate; pinnules pinnatipartite 1/2 or more to base; apex of pinnule lobe not falcate, rounded or rounded-truncate; perispore with few verrucose projections.   75 D. wheeleri
76 (71) Scales concolored, margin not black.   76 D. nipponicum
+ Scales ?bicolored, with black narrow margin conspicuous or inconspicuous   (77)
77 (76) Stipe, rachis, and costa dark brown, with dingy glandular hairs; sori shortly linear, inframedial from base upward to half of veinlets; huge plants with erect rhizomes ca. 70 cm tall, ca. 20 cm in diam..   77 D. axillare
+ Stipe (except base), rachis, and costa stramineous or green-stramineous, without dingy glandular hairs; sori mostly medial or inframedial from base to more than 2/3 of veinlet margin, or nearly as long as veinlet   (78)
78 (77) Rhizome usually ascending or erect, rarely creeping   (79)
+ Rhizome creeping   (82)
79 (78) Sori oblong, medial.   78 D. kappanense
+ Sori linear, mostly from base or near base to more than 2/3 of veinlet length, few sori short, brown, medial   (80)
80 (79) Stipe and rachis with sparse, fleshy, green, aculeate, verrucose processes; rhizome variable, erect, creeping; lamina thinly papery.   79 D. nanchuanicum
+ Stipe and rachis glabrous; lamina herbaceous   (81)
81 (80) Rhizome creeping to ascending; scales brown, lanceolate; pinnules broadly lanceolate or lanceolate, base asymmetrical, basal acroscopic pinnule lobes usually small; lamina abaxially with sparse light brown scales, without glands.   80 D. calogrammoides
+ Rhizome ascending to erect; scales mostly dark brown, linear- lanceolate, apex filiform or caudate; pinnules mostly deltoid- lanceolate, few lanceolate, symmetrical; costule and midrib with light brown or gray-white thin glands, rachis and costae abaxially with sparse linear, small scales.   81 D. viridissimum
82 (78) Costa and costule abaxially with prominent green fleshy aculeate processes; sori mostly as long as veinlets; black margin of scales incomplete.   82 D. calogrammum
+ Costa and costule without green fleshy aculeate processes   (83)
83 (82) Lamina thickly papery; pinnules broadly lanceolate or oblong-lanceolate; pinnule lobes of lower pinnae (at least those of lower pinnules) acute at apex, only pinnule lobes of upper pinnae or lobes of upper pinnules of middle or lower pinnae rounded at apex.   83 D. succulentum
+ Lamina herbaceous or thinly herbaceous; pinnule lobes rounded or truncate at apex   (84)
84 (83) Evergreen; pinnules mostly broadly deltoid-lanceolate, upper pinnules rarely lanceolate; sori thin, long, mostly as long as veinlets; stipe usually with sparse verrucose processes and slightly rough.   84 D. maximum
+ Summer-green; pinnules broadly lanceolate or oblong-lanceolate; sori mostly shortly linear, from base upward to 2/3 of veinlet length, distal part of lobes not soriferous; stipe without verrucose processes, smooth   (85)
85 (84) Pinnules lanceolate or oblong-lanceolate, glandular; sori elongate, from base to more than 2/3 of veinlet length.   85 D. giganteum
+ Pinnules oblong-lanceolate, glabrous; sori shortly linear, medial-inframedial.   86 D. latipinnulum

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