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Dryopteridaceae This PDF version does not have an ISBN or ISSN and is not therefore effectively published (Melbourne Code, Art. 29.1). The printed version, however, was effectively published on 6 June 2013. Zhang, L. B., S. G. Wu, J. Y. Xiang, F. W. Xing, H. He, F. G. Wang, S. G. Lu, S. Y. Dong, D. S. Barrington, K. Iwatsuki, M. J. M. Christenhusz, J. T. Mickel, M. Kato & M. G. Gilbert. 2013. Dryopteridaceae. Pp. 541–724 in Z. Y. Wu, P. H. Raven & D. Y. Hong, eds., Flora of China, Vol. 2–3 (Pteridophytes). Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. DRYOPTERIDACEAE 鳞毛蕨科 lin mao jue ke Zhang Libing (张丽兵)1, Wu Sugong (武素功)2, Xiang Jianying (向建英)2, Xing Fuwu (邢福武)3, He Hai (何海)4, Wang Faguo (王发国)3, Lu Shugang (陆树刚)5, Dong Shiyong (董仕勇)3; David S. Barrington6, Kunio Iwatsuki7, Maarten J. M. Christenhusz8, John T. Mickel9, Masahiro Kato10, Michael G. Gilbert11 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 articu- late 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, achloro- phyllous, 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, Nothopera- nema, 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, Steno- lepia 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 morphol- ogy 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. Key to subfamilies and genera 1a. Fronds weakly to strongly dimorphic; sporangia nearly evenly covering fertile parts of fronds abaxially (2. Subfam. Elaphoglossoideae). 2a. Stems climbing. 3a. Venation of sterile fronds anastomosing; lamina catadromous (or isodromous) toward apex ................... 9. Lomagramma 3b. Venation of sterile fronds free; lamina often anadromous toward apex ................................................... 10. Teratophyllum 2b. Stems (rhizomes) erect or creeping. 4a. Stipe base not articulate; lamina pinnate or rarely bipinnate, if simple then with reticulate venation; venation often reticulate .......................................................................................................................................... 7. Bolbitis 1 Missouri Botanical Garden, P.O. Box 299, Saint Louis, Missouri 63166-0299, U.S.A.; Chengdu Institute of Biology, Chinese Academy of Sciences, P.O. Box 416, Chengdu, Sichuan 610041, People’s Republic of China. 2 Kunming Institute of Botany, Chinese Academy of Sciences, 132 Lanhei Road, Heilongtan, Kunming, Yunnan 650204, People’s Republic of China. (Wu Songong died on 9 March 2013.) 3 South China Botanical Garden, Chinese Academy of Sciences, 723 Xingke Road, Tianhe District, Guangzhou, Guangdong 510650, People’s Republic of China. 4 Biological Herbarium III, Chongqing Normal University, Yifu Experimental Building, Huxi Campus, Shapingba District, Chongqing 401331, People’s Republic of China. 5 Herbarium, College of Life Science, Yunnan University, 2 Cuihu North, Kunming, Yunnan 650091, People’s Republic of China. 6 Pringle Herbarium, University of Vermont, Torrey Hall, 27 Colchester Avenue, Burlington, Vermont 05405, U.S.A. 7 815-29 Kamoshida, Aoba-ku, Yokohama 227-0033, Japan. 8 Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3SD, United Kingdom. 9 Institute of Systematic Botany, New York Botanical Garden, Bronx, New York 10458-5126, U.S.A. 10 Department of Botany, National Museum of Nature and Science, Amakubo 4-1-1, Tsukuba 305-0005, Japan. 11 Missouri Botanical Garden, c/o Herbarium, Library, Art and Archives, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, United Kingdom. 541 542 DRYOPTERIDACEAE 4b. Stipe base articulate (sometimes obscurely so); lamina simple; venation usually free ............................ 8. Elaphoglossum 1b. 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). 5a. Fronds often with ctenitoid hairs; costae and costules adaxially convex, not grooved. 6a. 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 ................................................................................................................................... 5. Lastreopsis 6b. 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. 7a. 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 ....................................... 2. Ctenitis 7b. 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) ............................ 4. Dryopteris 5b. Fronds without ctenitoid hairs; costae and costules adaxially grooved. 8a. 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. 9a. 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) ................................. 4. Dryopteris 9b. 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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