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6. Polystichum Roth, Tent. Fl. Germ. 3: 31, 69. 1799.

耳蕨属 er jue shu

Authors: Li-Bing Zhang & David S. Barrington

Acropelta Nakai; Cyrtomidictyum Ching; Hemesteum H. Léveillé (1915), not Newman (1851); Plecosorus Fée.

Plants perennial, terrestrial, evergreen or summer-green. Rhizome short, erect or ascending, together with basal stipe often covered with scales; scales linear to ovate, rarely with hairs. Fronds caespitose, monomorphic or rarely nearly dimorphic; stipe stramineous to purplish brown, canaliculate adaxially, scaly; distal stipe scales sparser, similar to but smaller than basal stipe scales. Lamina linear-lanceolate, lanceolate, ovate-lanceolate, lorate, or oblong, pinnate, bipinnatifid, or bipinnate, rarely finely divided and tripinnatifid or tetrapinnatifid; frond texture thinly papery, papery, or leathery, with lanceolate, subulate, linear, or ovate microscales abaxially and sometimes also adaxially, acroscopic base often auriculate, rarely auricles indistinct; rachis sulcate adaxially, with or without proliferous bulbils, rarely proliferous bulbils borne terminally on a prolonged flagelliform rachis; venation pinnate, free or rarely anastomosing to form 1 or 2 rows of areoles. Sori orbicular, terminal on veins of pinnae, sometimes abaxial or nearly terminal on veins, indusiate or rarely exindusiate; indusia orbicular, peltate, membranous, entire, erose, or irregularly toothed.

About 500 species: common in the temperate regions and subtropical lowlands and montane to alpine areas in the N Hemisphere, mostly in S and SW China, Himalaya, Japan, and Vietnam; a rich diversity of Polystichum also found in Central and South America; ca. 15 species in North America north of Mexico, 16 in Africa, eight in Madagascar and the Mascarenes, three in Macronesia, four in Europe, 12 in Australasia, and a few in New Guinea and the Pacific islands; 208 species (139 endemic) in two subgenera in China.

Chinese species can be classified into two subgenera and 23 sections.

The following taxa are excluded from the present treatment, pending further research: Aspidium aculeatum (Linnaeus) Swarz var. veitchii Christ (Bull. Herb. Boissier, sér. 2, 3: 513. 1903), described from W Hubei, A. aculeatum var. yunnanense Christ (Bull. Herb. Boissier 6: 964. 1898), described from Yunnan, and A. braunii Spenner var. clarkei Christ (Bull. Herb. Boissier 7: 15. 1899 ["Clarkii"]), described from Yunnan.


Key to sections

1 Plants evergreen, rarely summer-green; lamina 1-pinnate; if lamina bipinnatifid to tetrapinnatifid then pinnae finely dissected (P. sect. Sphaenopolystichum) or rachis scales dark brown, ovate, and appressed (P. excelsius, P. tonkinense); venation free or rarely anastomosing to form 1 or 2 rows of areoles (P. balansae, P. fraxinellum, P. hookerianum, P. lepidocaulon, P. ×rupestris, P. trapezoideum, P. uniseriale) (P. subg. Haplopolystichum)   (2)
+ Plants evergreen or summer-green; lamina bipinnate or bipinnatifid; if lamina 1-pinnate then microscales broad-type (P. sect. Hecatoptera, P. sect. Stenopolystichum; excl. P. sect. Mastigopteris) but rachis not prolonged at apex (excl. P. sect. Cyrtomiopsis) and rachis scales not ovate-lanceolate (excl. P. loratum); or if 1-pinnate then sori abaxial on veinlets (P. chingiae, P. nepalense); or if 1-pinnate then pinnae stiff and with open hard spinules on margin (P. delavayi, P. lonchitis); or if 1-pinnate then pinnae with long fine microscales and rachis without bulbils (P. fimbriatum, P. muscicola, P. subfimbriatum, P. zayuense, some species of P. sect. Sorolepidium), or pinnae ovate-deltoid and with broadly lanceolate microscales (P. duthiei, P. glaciale); venation free (P. subg. Polystichum)   (10)
       
2 (1) Rachis with proliferous bulbils   (3)
+ Rachis without proliferous bulbils   (5)
       
3 (2) Microscales broad-type; rachis prolonged at apex; sori exindusiate.   15 P. sect. Cyrtomiopsis
+ Microscales narrow-type; rachis prolonged or not prolonged at apex; sori indusiate   (4)
       
4 (3) Indusia entire; rachis prolonged at apex.   16 P. sect. Mastigopteris
+ Indusia erose; rachis prolonged (P. erosum) or not prolonged at apex.   17 P. sect. Basigemmifera
       
5 (2) Lamina with extraordinarily prolonged and 1-pinnate basal pinnae.   18 P. sect. Crucifilix
+ Lamina without extraordinarily prolonged basal pinnae   (6)
       
6 (5) Lamina bipinnatifid to tetrapinnatifid, rarely 1-pinnate; pinnae finely dissected.   19 P. sect. Sphaenopolystichum
+ Lamina 1-pinnate; pinnae not finely dissected   (7)
       
7 (6) Lamina leathery; if lamina papery then pinnae long spinulose on margin (P. formosanum); pinnae falcate-lanceolate, length more than 3 ?width, middle pinnae 0.9-2 cm wide; if middle pinnae 0.6-0.9 cm wide then pinnae obliquely oblong, up to 3.2 cm (P. minimum) or pinnae long spinulose on margin (P. formosanum); venation free or anastomosing to form 1 or 2 rows of areoles   (8)
+ Lamina papery or thinly papery; pinnae oblong or lanceolate, length normally 1-2 ?width, middle pinnae much narrower than 1 cm except those of P. acutidens; venation free   (9)
       
8 (7) Sori in 1 row on each side of midrib; pinnae symmetrical or nearly symmetrical at base; if pinnae asymmetrical at base then pinnae obliquely oblong, up to 3.2 ?0.9-2 cm (P. minimum); venation free; if venation anastomosing to form 1 or 2 rows of areoles along midrib then pinnae up to 10 pairs (P. fraxinellum).   20 P. sect. Cyrtogonellum
+ Sori in 2 or more rows on each side of midrib; if sori in 1 row on each side of midrib then sori close to pinna margin (P. falcatilobum, P. formosanum); pinnae asymmetrical at base; venation anastomosing to form 1 or 2 rows of areoles; if venation free then sori in 1 row on each side of midrib.   21 P. sect. Adenolepia
       
9 (7) Rachis scales ovate to lanceolate.   22 P. sect. Platylepia
+ Rachis scales linear, subulate or narrowly lanceolate.   23 P. sect. Haplopolystichum
       
10 (1) Lamina stiff, leathery or nearly leathery, often shiny adaxially; pinnae dentate and with hard spinules at apex and often also on margin   (11)
+ Lamina not stiff, papery; if lamina leathery then pinnae without hard spinules on margin (P. fimbriatum, P. subfimbriatum), dull or shiny adaxially; pinnae without hard spinules at apex or margin   (12)
       
11 (10) Lamina 1-pinnate; sori abaxial on veinlets.   1 P. sect. Polystichum
+ Lamina bipinnate, rarely 1-pinnate (P. delavayi) or tripinnatifid; sori terminal on veinlets.   2 P. sect. Xiphopolystichum
       
12 (10) Lamina 1-pinnate; rachis without proliferous bulbils; pinnae densely regularly long spinulose on margin.   3 P. sect. Hecatoptera
+ Lamina 1-pinnate or bipinnate; rachis with or without proliferous bulbils; if lamina 1-pinnate then pinnae shortly spinulose or entire on margin   (13)
       
13 (12) Lamina 1-pinnate; rachis with proliferous bulbils.   4 P. sect. Stenopolystichum
+ Lamina bipinnate or 1-pinnate; rachis without or with proliferous bulbils; if lamina 1-pinnate then rachis without proliferous bulbils   (14)
       
14 (13) Sori abaxial on veinlets; lamina green   (15)
+ Sori terminal on veinlets; if sori abaxial on veinlets then lamina dark green (P. biaristatum, P. mucronifolium, P. parvifoliolatum)   (18)
       
15 (14) Fronds up to 1 m or longer; lamina bipinnate.   8 P. sect. Neopolystichum
+ Fronds shorter than 50 cm; lamina 1-pinnate or bipinnatifid; if lamina bipinnatifid then pinnae and pinnules cartilaginous at margin   (16)
       
16 (15) Lamina 1-pinnate or bipinnatifid; pinnae and/or pinnules cartilaginous at margin; sori in 1 row on each side of midrib.   5 P. sect. Achroloma
+ Lamina 1-pinnate; pinnae not cartilaginous at margin; sori in 2 or 3 rows on each side of midrib   (17)
       
17 (16) Auricles well developed and deltoid, sharply pointed at apex.   6 P. sect. Chingiarum
+ Auricles less developed and rounded.   7 P. sect. Crinigera
       
18 (14) Lamina 1-pinnate; fronds leathery   (19)
+ Lamina bipinnate or bipinnatifid; fronds papery or thinly leathery; if lamina 1-pinnate then lamina papery   (20)
       
19 (18) Pinnae oblong; auricles sharply pointed at apex.   9 P. sect. Fimbriata
+ Pinnae ovate; auricles obtuse at apex.   10 P. sect. Subfimbriata
       
20 (18) Fronds shorter than 40(-60) cm, thinly papery; lamina 1-pinnate or shallowly to deeply pinnatifid.   11 P. sect. Micropolystichum
+ Fronds often longer than (40-)60 cm and up to 1 m or longer, papery; lamina bipinnate   (21)
       
21 (20) Plants summer-green; lamina 1-pinnate to bipinnatisect; microscales fibrillose or lanceolate, longer than 0.8 mm.   12 P. sect. Sorolepidium
+ Plants evergreen or summer-green; lamina bipinnate; microscales linear, shorter than. 5(-0.6) mm; if microscales fibrillose and up to 1 mm then pinnules exauriculate   (22)
       
22 (21) Plants evergreen; lamina dark green, shiny adaxially, up to 100 cm or longer; rachis with or without proliferous bulbils; pinnules lanceolate.   13 P. sect. Macropolystichum
+ Plants evergreen or summer-green; lamina green, normally shorter than 80 cm, dull adaxially; rachis without proliferous bulbils; pinnules ovate-lanceolate   14 P. sect. Hypopeltis

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