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

1. Deparia Hooker & Greville, Icon. Filic. 2: t. 154. 1829.

对囊蕨属 dui nang jue shu

Authors: He Zhaorong, Zhongren Wang & Masahiro Kato

Athyriopsis Ching; Dictyodroma Ching; Dryoathyrium Ching; Lunathyrium Koidzumi; Parathyrium Holttum; Triblemma Ching.

Plants terrestrial, medium-sized. Rhizomes moderately stout, long creeping, ascending or suberect, with black or brown, lanceolate, entire or subentire scales. Fronds distant or approximate; stipe long, with brown, ovate to lanceolate scales at base; lamina pinnate or 2-pinnate, lanceolate, linear-lanceolate, oblong, or ovate-oblong, acuminate; ultimate pinnules pinnatipartite; pinnae or pinnules alternate, sessile, equilateral, rounded-cuneate at base, pinnatifid to pinnatipartite, segments oblong or rectangular, pinnules or segments adnate to costae or costules by narrow wings; rachis, costae, and costules grooved adaxially, grooves of costae or costules interrupted and not merging into grooves of rachis or costae; usually with brown or dark brown, vermiculate glandular hairs consisting of 1-3(or 4) rows of cells on costae, costules, and midribs. Veins free, lateral veins simple or forked. Lamina herbaceous, papery, or subleathery when dried. Sori orbicular, oblong, J-shaped, or horseshoe-shaped, abaxial or lateral, medial; indusia same in shape, membranous, entire, erose, lacerate, or ciliate, persistent. Spores bilateral, perispore with folds or spines, baculate ornamentation. x = 40.

About 70 species: tropical and temperate zones of Asia, tropical Africa, and Madagascar, extending to Japan, Korea, E Russia, and to NW Himalaya; 53 species (31 endemic) in China.

The following regions comprise the center of distribution in China: Qin Ling, W Hubei, Sichuan, and NE Yunnan.

The following taxon is excluded from the present treatment, pending further research: Lunathyrium ebeneostipes Ching & Z. Y. Liu (Bull. Bot. Res., Harbin 3(4): 14. 1983), described from Sichuan (Nanchuan).

Putative Hybrids

From abortive spores and morphological features of specimens, many hybrids can be considered. Here only two hybrids that have been published as “species” are listed.

Deparia ×kanghsienense (Ching & Y. P. Hsu) Z. R. He, comb. nov.

康县对囊蕨 kang xian dui nang jue

Basionym: Lunathyrium ×kanghsienense Ching & Y. P. Hsu, Fl. Tsinling. 2: 212. 1974, pro sp.

The type of this hybrid is morphologically intermediate between Deparia liangshanensis var. sericea and D. sichuanensis. It is similar to D. liangshanensis in the multicellular long hairs on the surface of indusia but differs by the pinna segments without long multicellular hairs at margin, at most occasionally with few short hairs between teeth. The hairs on pinna segments are much denser than in D. sichuanensis. The spores are mostly abortive.

● 1300-1400 m. Gansu (Kangxian).

Deparia ×nanchuanense (Ching & Z. Y. Liu), Z. R. He, comb. nov.

南川对囊蕨 nan chuan dui nang jue

Basionym: Lunathyrium ×nanchuanense Ching & Z. Y. Liu, Bull. Bot. Res., Harbin 3(4): 15. 1983, pro sp.

This hybrid is morphologically intermediate between Deparia shennongensis and D. vegetior. It is similar to D. vegetior in the hairs on the abaxial surface of rachis and costae very thin, short, and sparse, and multicellular hairs absent at sinuses between pinna segments, and multicellular hairs on the adaxial surface of veins in fertile pinnae very short. It is also similar to D. shennongensis in the lower pinnae gradually shortened and spore surface with rugate folds or baculate processes. Some spores in each sporangium are abortive.

● Forests; 1600-1700 m. Chongqing (Jinfo Shan).

There are other formerly undescribed species, e.g., “Lunathyrium subimbricatum” (Ching in Y. L. Zhang et al., Sporae Pterid. Sin. 233, t. 45, f. 21, 25. 1976; 栉比蛾眉蕨 jie bi e mei jue) in broad-leaved forests, rock crevices, watersides, 2700-2800 m, W Yunnan (Yongping); and “Lunathyrium heterocarpum” (Ching in Y. L. Zhang et al., Sporae Pterid. Sin. 232, t. 46, f. 10, 15. 1976; 杂果蛾眉蕨 za guo e mei jue) in forests on mountain slopes, ca. 3300 m, Sichuan (Li Xian). The former is similar to Deparia sichuanensis, but the plant is larger with basal pinna segments prominently broadened. It is also similar to D. wilsonii. The spores are completely abortive. The latter hybrid is intermediate between D. dolosa and D. shennongensis in its pinnae to 30 pairs or more, pinna segments to 20 pairs or more, and rachis and costae with rather dense multicellular long hairs abaxially. The spores are partly abortive, and “good” spores are variable in size.

1 Stipe base not swollen, without pneumatophore   (2)
+ Stipe base swollen, usually winged with pneumatophore   (23)
2 (1) Veins anastomosing   (3)
+ Veins free   (6)
3 (2) Free fertile pinnae repand or pinnatilobate at margin; lateral veins obvious, costae with sparse hairs   (4)
+ Free fertile pinnae entire, undulate, or shallowly lobed; lateral veins not obvious, costae subglabrous   (5)
4 (3) Free pinnae of fertile frond 1-5 pairs, margin usually repand, occasionally crenate or pinnatilobate; pinna lobes flat, broad, blunt, oval, symmetrical.   33 D. heterophlebia
+ Free pinnae of fertile frond 6-8 pairs, margin shallowly lobed; pinna lobes deltoid-oblong, asymmetrical, subfalcate, apex obtuse.   34 D. yunnanensis
5 (3) Fertile lamina deeply pinnatilobate or 1-pinnate with 1 or 2(or 3) free basal pairs of pinnae; pinnae oblong-lanceolate, base rounded.   35 D. formosana
+ Fertile lamina 1-pinnate with 3-6 pairs of free lateral pinnae; lower pinnae usually broadly lanceolate, sometimes oblong-lanceolate, base subcordate.   36 D. hainanensis
6 (2) Lamina simple or pinnatilobate; rachis glabrous, costa abaxially glabrous; sori elongate   (7)
+ Lamina 1-3-pinnate; rachis hairy, costa abaxially hairy; sori variable (elongate, horseshoe-shaped, J-shaped, or orbicular-reniform)   (8)
7 (6) Lamina simple, margin entire or repand.   37 D. lancea
+ Lamina pinnatilobate.   38 D. tomitaroana
8 (6) Rhizome ascending or erect; fronds caespitose   (9)
+ Rhizome slender, creeping; fronds distant to approximate   (11)
9 (8) Rhizome erect; indusia lacerate, ciliate at margin, perispore surface with sparse auriculate and aculeate projections.   39 D. omeiensis
+ Rhizome ascending; indusia subentire, slightly denticulate, or erose   (10)
10 (9) Lamina ovate-oblong; indusia erose at margin; perispore surface with dense unevenly long aculeate projections.   40 D. concinna
+ Lamina narrowly ovate or ovate; indusia subentire or slightly denticulate; perispore surface with dense clavate and thickly aculeate projections.   41 D. dickasonii
11 (8) Lamina narrowly lanceolate, lanceolate, broadly lanceolate, or narrowly deltoid; pinnae rounded and acute at apex   (12)
+ Lamina ovate, oblong, deltoid, broadly lanceolate, or broadly oblong-lanceolate, acuminate to long acuminate at apex, rarely acute   (16)
12 (11) Lamina herbaceous, rachis and costae with often many curly long nodose hairs on both surfaces; free pinnae 1 or 2(or 3) pairs; indusia with short, nodose hairs or glabrous, margin lacerate, ciliate, usually flat, rarely incurved; perispore with dense and apically rounded, aculeate projections.   42 D. petersenii
+ Lamina thinly herbaceous or submembranous, with sparsely nodose hairs; free pinnae often more than 5 pairs, rarely 2 or 3 pairs   (13)
13 (12) Lamina narrowly lanceolate, lanceolate, or broadly lanceolate, 3-5 ?as long as wide; indusial margin usually erose, few lacerate   (14)
+ Lamina narrowly deltoid, 2-3 ?as long as wide; indusial margin lacerate, ciliate, flat when young   (15)
14 (13) Lamina narrowly or broadly lanceolate, 4-5 ?as long as wide, less than 5 cm wide; indusial margin mostly flat when young, rarely incurved; perispore with verrucose and fossulate (muriform) ornamentation.   43 D. conilii
+ Lamina broadly lanceolate, ca. 3 ?as long as wide, 8-11 cm wide; indusial margin incurved when young; surface of perispore with irregular tuberculate ornamentation.   44 D. lushanensis
15 (13) Free pinnae more than 5 pairs, basal pinnae prominently longer than above; indusial surface with ?tiny nodose hairs.   45 D. pseudoconilii
+ Free pinnae 2 or 3 pairs, basal 1 pair often slightly reduced or nearly as long as second one; indusial surface glabrous.   46 D. pachyphylla
16 (11) Lamina broadly lanceolate or oblong-lanceolate, 2-4 ?as long as wide, less than 12 cm wide, apex acuminate or acute; margin of indusia shallowly erose or subentire, involute when young   (17)
+ Lamina ovate, oblong, deltoid, or narrowly deltoid, 2 ?or less as long as wide, apex acuminate to long acuminate; margin of indusia deeply or shallowly lacerate, often erose when old, incurved or flat when young   (18)
17 (16) Lamina broadly lanceolate, basal pinnae not reduced; sori subcostular, perispore surface with irregular auriculate ornamentation.   47 D. shandongensis
+ Lamina oblong or broadly lanceolate, 2 or 3 basal pairs of pinnae gradually reduced; basal pair of pinnae oblique, ovate, ca. 1/2 ?as long as middle pinnae; sori medial, perispore surface densely long aculeate and with long clavate ornamentation.   48 D. abbreviata
18 (16) Free pinnae mostly ascending at 60?angle, base broadly cuneate or cuneate, pinna lobes prominently ascending, with sparse nodose hairs on both surfaces of lamina; surface of indusia glabrous.   49 D. japonica
+ Free pinnae usually ascending at angles more than 70?or spreading, pinna lobes spreading or oblique at angles more than 50?(often 60?70?; rachis and costae abaxially usually with prominently thick, long, nodose hairs; adaxial surface of pinna with slender pointed short nodose hairs; surface of indusia hairy or glabrous   (19)
19 (18) Stipe and rachis with dense pale brown, transparent or translucent, lanceolate to linear, thickly clathrate membranous scales and thick nodose hairs 2-3 mm   (20)
+ Stipe and rachis below often with nodose hairs ca. 1.5 mm and sparsely scaly   (21)
20 (19) Pinna lobes entire or undulate at margin, at most shallowly crenate, rachis with dense thick long nodose hairs on abaxial side, mixed with 2- or 3-cells-wide, thickly clathrate, long linear scales; indusial surface hairy, margin flat when young, perispore surface densely thick clavate and with aculeate ornamentation.   50 D. kiusiana
+ Pinna lobes crenate at margin; basal and medial parts of rachis dense with 2- or 3-cells-wide, thickly clathrate, long linear scales, mixed with few thick, long nodose hairs, upper part of rachis with sparse scales and many nodose hairs; indusial surface glabrous, margin mostly flat when young, partly incurved, perispore surface with dense mostly truncate and clavate ornamentation.   51 D. jinfoshanensis
21 (19) Pinnae below middle broadly cuneate or cuneate; pinna lobes often oblong or ligulate-oblong, apex acute or truncate, oblique, rarely rounded.   42 D. petersenii
+ Pinnae below middle shallowly cordiform or truncate at base; pinna lobes ligulate-oblong or oblique, falcate oblong, apex rounded   (22)
22 (21) Pinna lobes usually entire or repand, sometimes crenate; indusial surface hairy, margin lacerate, incurved when young; perispore surface with dense thick uniformly slender clavate ornamentation.   52 D. dimorphophyllum
+ Pinna lobes usually serrate, or finely repand; indusial surface usually glabrous, margin lacerate, usually flat when young, rarely incurved; perispore surface with most apically truncate clavate, sometimes narrow, flakelike projections.   53 D. longipes
23 (1) Lowest pinnule not auricled and slightly reduced; sori linear, J-shaped, U-shaped, or orbicular-reniform, not or rarely diplazioid   (24)
+ Lowest pinnule auricled and slightly to hardly reduced; sori linear, J-shaped, diplazioid, or U-shaped   (36)
24 (23) Sori oblong or shortly linear, sometimes curved; indusia oblong, lunate, J-shaped, or horseshoe-shaped   (25)
+ Sori mostly orbicular or elliptic; indusia orbicular-reniform, horseshoe-shaped, J-shaped, or lunate   (30)
25 (24) Lamina 3-pinnate; pinnules stalked, pinnatilobate to pinnatipartite at margin, lateral veins pinnate.   1 D. pterorachis
+ Lamina pinnate-pinnatifid; pinnules entire or toothed at margin, lateral veins simple or forked   (26)
26 (25) Lamina ovate, pinnae stalked; stipe and rachis with dense multicellular hairs; spore surface with long spines.   2 D. erecta
+ Lamina oblong, pinnae sessile; stipe and rachis with sparse vermiculate hairs consisting of 2 or 3 rows of cells; spore surface with folded or rugate processes   (27)
27 (26) Segments entire (occasionally repand), lateral veins simple; sori mostly shortly linear (sometimes slightly curved), occasionally J-shaped.   3 D. chinensis
+ Segments toothed, lateral veins 2- or 3(or 4)-forked; sori horseshoe-shaped, J-shaped, oblong, or elliptic   (28)
28 (27) Lamina with sparse brown spinelike thick hairs adaxially; lateral veins usually forked.   4 D. setigera
+ Lamina subglabrate, without spinelike thick hairs adaxially; lateral veins usually 2-4-forked   (29)
29 (28) Rhizomes ascending or suberect; sori usually narrowly oblong, lunate, or J-shaped.   5 D. coreana
+ Rhizomes shortly creeping; sori shortly oblong, J-shaped, or horseshoe-shaped.   6 D. henryi
30 (24) Lamina 2-pinnate-pinnatifid with pinnules free or connected by costal wing   (31)
+ Lamina 2-pinnatipartite with pinnules adnate to costa   (33)
31 (30) Pinnules free, shortly stalked.   13 D. boryana
+ Pinnules ?connected with costa   (32)
32 (31) Lamina thickly herbaceous, pinnules subquadrangular at base, pinnatifid 1/2 or less to costule, segments entire or repand-toothed.   11 D. okuboana
+ Lamina thinly herbaceous, pinnules broadly cuneate at base, pinnatifid more than 2/3 to costule, segments crenate.   12 D. viridifrons
33 (30) Sori elliptic or orbicular-elliptic, indusia lunate or horseshoe-shaped.   7 D. confusa
+ Sori orbicular or suborbicular, indusia orbicular-reniform, occasionally horseshoe-shaped   (34)
34 (33) Stipe and rachis with sparse dark brown ovate-lanceolate scales; segments crenate; veins forked to 3(or 4)-forked.   8 D. stenopterum
+ Stipe and rachis clothed with very many blackish brown, shiny, broadly lanceolate scales; segments entire or toothed; lateral veins forked, occasionally simple or 3-forked   (35)
35 (34) Pinnae narrowly elliptic or elliptic, pinnatisect nearly to costae; pinna segments unevenly long, basiscopic segments longer than acroscopic segments, often falcate.   9 D. falcatipinnula
+ Pinnae lanceolate, pinnatifid, segments even, oblong.   10 D. unifurcata
36 (23) Rhizomes ascending or shortly creeping; lamina with only 3 pairs of lower pinnae slightly shortened, basal pair of pinnae usually more than 3 cm; multicellular short hairs present at sinus between segments.   14 D. pycnosora
+ Rhizomes erect; lamina with more pairs of lower pinnae gradually shortened or only basal 2 or 3 pairs slightly shortened; multicellular short hairs absent or present at sinus between segments; perispore surface with rugose, rugate, saccate, papillate, baculate processes, or ridged folds   (37)
37 (36) Multicellular hairs ?present at sinus between segments   (38)
+ Multicellular hairs absent at sinus between segments   (42)
38 (37) Lamina with only few pairs of lower pinnae slightly shortened, basal pair of pinnae never shortened as auriculate, usually more than 2 cm, stipes usually more than 10-15 cm   (39)
+ Lamina with numerous pairs of lower pinnae gradually shortened toward base, basal pair of pinnae no more than 1-2 cm, often auriculate, stipes usually within 10-15 cm   (40)
39 (38) Indusia with short glandular hairs on back.   15 D. auriculata
+ Indusia without or with short glandular hairs only on lower part of back.   16 D. giraldii
40 (38) Multicellular hairs sparsely present on surfaces of segments.   19 D. sichuanensis
+ Multicellular long hairs densely present on surfaces of segments   (41)
41 (40) Spore surface with few discontinuous low folds or rugose folds; rachis and costae with dense multicellular long hairs and scale hairs; hairs absent at margin of segments.   17 D. hirtirachis
+ Spore surface with rugate, saccate folds, or papillate, verrucose processes; rachis and costae only with sparse multicellular hairs; segments with multicellular long hairs at margin.   18 D. liangshanensis
42 (37) Perispore surface with continuous low folds or discontinuous rugose folds   (43)
+ Perispore surface with rugate, saccate folds or papillate, verrucose, or baculate processes   (46)
43 (42) Numerous pairs of lower pinnae gradually shortened toward base, basal pair of pinnae no more than 2 cm, often auriculate   (44)
+ Only 2 or 3 pairs of lower pinnae slightly shortened toward base, basal pair of pinnae usually more than 3 cm   (45)
44 (43) Fertile fronds only ca. 30 cm; segments thinner, less than 2 mm wide, subentire at margin, pinnae and segments approximate; multicellular hairs thicker and longer on abaxial surface of rachis and costae.   20 D. sikkimensis
+ Fertile fronds more than 60 cm; segments more than 4 mm wide, serrulate at margin, pinnae and segments separate; multicellular hairs thin and short on abaxial surface of rachis and costae.   21 D. medogensis
45 (43) Multicellular hairs very short on adaxial surface of fertile pinnae, inconspicuous; hairs thin, short, and sparse on abaxial surface of rachis and costae; indusia subentire.   22 D. vegetior
+ Multicellular hairs visible on adaxial surface of fertile pinnae; hairs thick and long on abaxial surface of rachis and costae; indusia slightly erose or shortly ciliate at margin.   23 D. acuta
46 (42) Only few pairs of lower pinnae slightly shortened toward base, basal pair of pinnae usually more than 2 cm, never auriculate, stipes usually more than 10-15 cm   (47)
+ Numerous pairs of lower pinnae gradually shortened toward base, basal pair of pinnae no more than 1-2 cm, often auriculate, stipes usually 10-15 cm   (49)
47 (46) Spore surface with dense baculate processes or spines.   24 D. emeiensis
+ Spore surface with sparse rugate, saccate folds or papillate, verrucose processes   (48)
48 (47) Lower pinnae narrowly deltoid-ovate or ovate-lanceolate, prominently broadened toward base, their basal segments reflexed.   25 D. ludingensis
+ Lower pinnae lanceolate, not broadened toward base, their basal segments explanate   26 D. wilsonii
49 (46) Multicellular hairs thin, short, and sparse at abaxial surfaces of rachis and costae   (50)
+ Multicellular hairs thick, long, and rather dense on abaxial surfaces of rachis and costae   (51)
50 (49) Segments truncate at apex.   27 D. vermiformis
+ Segments obtuse-rounded or obtuse at apex.   28 D. shennongensis
51 (49) Segments truncate at apex.   29 D. truncata
+ Segments rounded or obtuse, occasionally rounded-truncate at apex   (52)
52 (51) Fertile fronds less than 30 cm; pinnae fewer than 12 pairs, less than 5 cm; segments 5-7 pairs.   30 D. brevipinna
+ Fertile fronds more than 30 cm; pinnae up to 20 pairs, more than 5 cm; segments more than 12 pairs   (53)
53 (52) Fertile fronds up to more than 1 m, lamina linear-oblanceolate, pinnae more than 25-30 pairs.   31 D. dolosa
+ Fertile fronds 30-70 cm, lamina oblong or oblanceolate, pinnae 20-25 pairs.   32 D. jiulungensis

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