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33. Dryopteridaceae
鳞毛蕨科 lin mao jue ke
Authors: Li-Bing Zhang, Sugong Wu, Xiang Jianying, Prof. Fuwu Xing, He Hai, Wang Faguo, Shugang Lu, Shiyong Dong, David S. Barrington, Kunio Iwatsuki, Maarten J. M. Christenhusz, John T. Mickel, Masahiro Kato & Michael G. Gilbert
Plants small to large, evergreen or deciduous, terrestrial, epilithic, hemiepiphytic, or epiphytic. Rhizomes erect, ascending, creeping, or sometimes climbing, dictyostelic (mostly radially), scaly; scales basally attached or very rarely peltate, often non-clathrate but sometimes clathrate, entire or dentate. Fronds caespitose or remote from one another, with segments anadromously or catadromously arranged, or sometimes anadromous at base and catadromous distally; stipe often not articulate but sometimes articulate at base, with 3 or more vascular bundles arranged in a semicircle or circle, scaly, sulcate adaxially, not hairy or sometimes hairy; lamina monomorphic or dimorphic, usually oblong, deltoid, pentagonal, lanceolate, ovate, or linear, 1-5-pinnate, or simple, rarely imparipinnate, scaly, glandular, hairy, or glabrous; if scaly then scales bullate or flat; glands absent or present; texture thinly papery, papery, or leathery. Rachises sulcate adaxially, with or without proliferous bulbils, rarely proliferous bulbils borne at a prolonged flagelliform rachis terminal; venation pinnate and free, or variously anastomosing to form 1 to multiple rows of areoles, with or without included veinlets. Fertile fronds conform or weakly to strongly difform with sterile fronds. Sporangia usually in orbicular sori; sori terminal, subterminal, or dorsal on veins, indusiate or rarely exindusiate; if indusiate then indusia orbicular or reniform or rarely ovoid, superior, lateral, or rarely inferior, sessile or rarely stalked, entire or toothed; sometimes sporangia nearly evenly covering fertile parts abaxially (not forming orbicular sori); sporangia with 3-rowed, short to long stalks. Spores monolete, achlorophyllous, with prominent perispore. x = 41.
About 25 genera and 2,100 species: nearly cosmopolitan, but highest diversity found in E Asia (Dryopteris, Polystichum) and the New World (Ctenitis, Elaphoglossum); ten genera and 493 species (260 endemic) in two subfamilies in China.
Eighty percent of the 2,100 species are in four genera (Ctenitis, Dryopteris, Elaphoglossum, and Polystichum). Subfamily Dryopteridoideae includes 17 genera: Adenoderris J. Smith, Arachniodes (incl. Leptorumohra, Lithostegia, Phanerophlebiopsis), Coveniella Tindale, Ctenitis (incl. Ataxipteris), Cyclodium C. Presl, Cyrtomium, Dryopolystichum Copeland, Dryopteris (incl. Acrophorus, Acrorumohra, Dryopsis, Nothoperanema, Peranema, Revwattsia D. L. Jones), Lastreopsis, Maxonia C. Christensen, Megalastrum Holttum, Phanerophlebia C. Presl, Polystichopsis (J. Smith) Holttum, Polystichum (incl. Cyrtogonellum, Cyrtomidictyum, Papuapteris Christensen, Plecosorus, Sorolepidium), Rumohra Raddi, Stenolepia Alderwerelt, and Stigmatopteris C. Christensen. Subfamily Elaphoglossoideae includes eight genera: Arthrobotrya C. Christensen, Bolbitis (incl. Egenolfia), Elaphoglossum (incl. Microstaphyla C. Presl, Peltapteris Link), Lomagramma, Mickelia R. C. Moran, Labiak & Sundue, Olfersia Raddi, Polybotrya Humbolt & Bonpland ex Willdenow, and Teratophyllum. These genera are placed in Dryopteridaceae on the basis of morphology and, for most, molecular studies as well. Wang Chuhao. 1999. Ctenitis, Lastreopsis, Bolbitidaceae, and Lomagramma. In: Wu Shiewhung, ed., Fl. Reipubl. Popularis Sin. 6(1): 2-37, 56-59, 104-124, 129-133; Wu Shiew-hung. 1999. Peranemataceae ["Peranemaceae"]. In: Wu Shiew-hung, ed., Fl. Reipubl. Popularis Sin. 4(2): 216-238; Wu Shiewhung. 1999. Elaphoglossaceae. In: Wu Shiewhung, ed., Fl. Reipubl. Popularis Sin. 6(1): 134-142; Hsieh Yingtang, Wu Sugong & Lu Shugang. 2000. Dryopteridaceae (1). In: Wu Sugong, ed., Fl. Reipubl. Popularis Sin. 5(1): 1-220; Kung Hsianshiu, Chu Weiming, He Zhaorong, Zhang Libing & Hsieh Yingtang. 2001. Dryopteridaceae (2) (excluding Cyclopeltis). In: Kung Hsianshiu, ed., Fl. Reipubl. Popularis Sin. 5(2): 1-228.
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Key to subfamilies and genera
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| 1 |
Fronds weakly to strongly dimorphic; sporangia nearly evenly covering fertile parts of fronds abaxially (2. Subfam. Elaphoglossoideae) |
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(2) |
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Fronds monomorphic; if dimorphic then rachis with proliferous bulbils borne at a prolonged flagelliform rachis terminal (Polystichum sect. Cyrtomiopsis); sporangia in orbicular sori (1. Subfam. Dryopteridoideae) |
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(5) |
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| 2 (1) |
Stems climbing |
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(3) |
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Stems (rhizomes) erect or creeping |
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(4) |
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| 3 (2) |
Venation of sterile fronds anastomosing; lamina catadromous (or isodromous) toward apex. |
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9 Lomagramma |
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Venation of sterile fronds free; lamina often anadromous toward apex. |
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10 Teratophyllum |
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| 4 (2) |
Stipe base not articulate; lamina pinnate or rarely bipinnate, if simple then with reticulate venation; venation often reticulate. |
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7 Bolbitis |
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Stipe base articulate (sometimes obscurely so); lamina simple; venation usually free. |
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8 Elaphoglossum |
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| 5 (1) |
Fronds often with ctenitoid hairs; costae and costules adaxially convex, not grooved |
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(6) |
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Fronds without ctenitoid hairs; costae and costules adaxially grooved |
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(8) |
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| 6 (5) |
Rhizomes often long and creeping or ascending; fronds often distant from one another; ratio of length to width of lamina often approximately 1:1; pinnule rachises and midribs often with yellow or reddish glands. |
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5 Lastreopsis |
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Rhizomes short and erect or ascending; fronds caespitose; ratio of length to width of lamina often clearly larger than 1:1; pinnule rachises and midribs often without glands |
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(7) |
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| 7 (6) |
Pinnules with veins indistinct on both adaxial and abaxial surfaces; sori dorsal on veinlets; scales ciliate on their margins, clathrate, and with nearly hexagonal and lustrous areoles. |
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2 Ctenitis |
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Pinnules with distinct veins on abaxial surfaces; sori terminal on veinlets, and marginal; scales entire and clathrate or not, but with long and dull areoles (Dryopteris sect. Dryopsis). |
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4 Dryopteris |
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| 8 (5) |
Sori if indusiate then indusia mostly reniform and lateral (attached to abaxial surface of fronds at notch of indusia), rarely orbicular and inferior (Dryopteris subg. Nothoperanema); if indusia ovoid then lamina finely 4- or 5-pinnate (Arachniodes superba); if sori exindusiate then ultimate segments of fronds not auriculate at acroscopic base (Dryopteris dehuaensis, D. gymnosora, D. liangkwangensis, D. polita, D. scottii, and D. shikokiana); venation free |
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(9) |
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Sori if indusiate then indusia orbicular and superior, peltate; if sori exindusiate then ultimate segments of fronds clearly auriculate at acroscopic base (Polystichum alcicorne, P. glaciale, P. grandifrons, P. gymnocarpium, P. longispinosum, P. loratum, and P. rufopaleaceum) or less auriculate but microscales long, i.e., longer than 1 mm (P. longipaleatum and P. nudisorum); venation free or anastomosing |
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(10) |
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| 9 (8) |
Rhizomes short and erect or ascending; lamina dull or lustrous adaxially; laminar segments arranged catadromously except secondary segments of basalmost pair of pinnae of bipinnate fronds; if laminar segments arranged anadromously then lamina (2 or)3- or 4-pinnate and ultimate pinnules of lamina asymmetrical at base (Dryopteris sect. Acrorumohra). |
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4 Dryopteris |
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Rhizomes often long and creeping or ascending; if rhizomes short then lamina finely 4- or 5-pinnate and indusia ovoid (Arachniodes superba); lamina lustrous adaxially; laminar segments arranged anadromously. |
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1 Arachniodes |
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| 10 (8) |
Lamina imparipinnate, apex with an apical pinna somewhat dissected at base, lamina rarely simple; venation always anastomosing to form 2 or more rows of areoles. |
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3 Cyrtomium |
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Lamina 1-3-pinnate, apex pinnatifid, without a clear apical pinna; venation mostly free, rarely anastomosing to form 1 or 2 rows of areoles (Polystichum balansae, P. fraxinellum, P. hookerianum, P. lepidocaulon, P. ?upestris, P. trapezoideum, and P. uniseriale). |
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6 Polystichum |
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List of lower taxa
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