Brief Summary

Comprehensive Description

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    The Dryopteridaceae are a family of leptosporangiate ferns in the order Polypodiales. They are known colloquially as the wood ferns. They comprise about 1700 species and have a cosmopolitan distribution. They may be terrestrial, epipetric, hemiepiphytic, or epiphytic. Many are cultivated as ornamental plants.[2] The largest genera are Elaphoglossum (600), Polystichum (260), Dryopteris (225), and Ctenitis (150). These four genera contain about 70% of the species.[3] Dryopteridaceae diverged from the other families in eupolypods I about 100 million years ago.[4]


    The rhizomes are often stout, creeping, ascending, or erect, and sometimes scandent or climbing, with nonclathrate scales at apices. Fronds are usually monomorphic, less often dimorphic, or sometimes scaly or glandular, but less commonly hairy. Petioles have numerous round, vascular bundles arranged in a ring, or rarely as few as three; the adaxial bundles are largest. Veins are pinnate or forking, free to variously anastomosing; the areoles occur with or without included veinlets; sori are usually round, acrostichoid (covering the entire abaxial surface of the lamina) in a few lineages; usually indusiate, or sometimes exindusiate. Indusia, when present, are round-reniform or peltate. Sporangia have three-rowed, short to long stalks; spores arereniform, monolete, perine or winged.[3]



    In 1990, Karl U. Kramer and coauthors defined the Dryopteridaceae broadly to include the present family, as well as the Woodsiaceae sensu lato, Onocleaceae, and most of Tectariaceae.[5] Molecular phylogenetic studies found Kramer's version of the Dryopteridaceae to be polyphyletic, and it was split up by Smith and others in 2006.[3] The inclusion of Didymochlaena, Hypodematium, and Leucostegia in the Dryopteridaceae is doubtful. If these three are excluded, then the family is strongly supported as monophyletic in cladistic analyses.[6] Some authors have already treated these genera as outside of the Dryopteridaceae.[1]

    In 2007, a phylogenetic study of DNA sequences showed that Pleocnemia should be transferred from the Tectariaceae to the Dryopteridaceae.[7] In 2010, in a paper on bolbitidoid ferns, Arthrobotrya was resurrected from Teratophyllum.[8] Later that year, Mickelia was described as a new genus.[9]

    Some species have been removed from the genus Oenotrichia because they do not belong there or even in the family Dennstaedtiaceae where Oenotrichia sensu stricto is placed. These species probably belong in the Dryopteridaceae, but have not yet been given a generic name.[6]

    In 2012, a phylogenetic study of Dryopteris and its relatives included Acrophorus, Acrorumohra, Diacalpe, Dryopsis, Nothoperanema, and Peranema within that genus.[10] The Flora of China treatment of the family, published in 2013, used phylogenetic results to sink Lithostegia and Phanerophlebiopsis into Arachniodes.[11]

    The Dryopteridaceae Herter, under the classification system of Christenhusz and Chase (2014), are submerged as subfamily Dryopteridoideae Link, one of eight subfamilies constituting family Polypodiaceae. This family corresponds to the clade Eupolypods I.[12]


    The PPG I classification divides the family into three subfamilies, listed below.[13]

    The following table shows the currently accepted Dryopteridaceae generic names and the corresponding synonyms.

    Accepted generic names[3] Synonyms[14][15] Arachniodes Blume 1828 Arthrobotrya J.Sm. 1875 Bolbitis Schott 1834
    • Anapausia C. Presl
    • Campium C. Presl
    • Cyrtogonium J.Sm.
    • Edanyoa Copel.
    • Egenolfia Schott 1836
    • Heteroneurum C.Presl
    • Jenkinsia Hook.
    • Poecilopteris C.Presl
    Ctenitis (C.Chr.) C.Chr. 1938
    • Atalopteris Maxon & C. Chr.
    • Ataxipteris Holttum 1984
    Cyclodium C.Presl 1836 Cyrtomium C.Presl 1836
    • Amblia C. Presl
    • Cyrtogonellum Ching 1938
    • Cyrtomidictyum Ching 1940
    Dryopteris Adans. 1763
    • Acrophorus C. Presl 1836[10]
    • Acrorumohra (H.Itô) H.Itô 1938[10]
    • Arthrobotrys (C.Presl) Lindl. 1846
    • Diacalpe Blume 1828
    • Dichasium (A.Braun) Fée 1852
    • Diclisodon T.Moore 1857
    • Dryopsis Holttum & P.J.Edwards 1986[10]
    • Filix Ség. 1754
    • Filix-mas Hill ex Farw. 1931
    • Lophodium Newman 1851
    • Nephrodium Marthe ex Michx. 1803
    • Nothoperanema (Tagawa) Ching 1966[10] – Island Lacefern[16]
    • Peranema D.Don 1825[10]
    • Pteris Gled. ex Scop. 1753
    • Pycnopteris T. Moore 1855
    • Revwattsia D.L.Jones 1998
    • Sphaeropteris R.Br. ex Wall. 1830 (non Bernh. 1801)[10]
    • Stenolepia Alderw. 1909
    Elaphoglossum Schott ex J. Sm. 1842
    • Aconiopteris C.Presl 1836
    • Dictyoglossum J.Sm. 1846
    • Hymenodium Fée 1845
    • Microstaphyla C.Presl 1851
    • Peltapteris Link 1841
    • Rhipidopteris Schott ex Fée 1845
    Lastreopsis Ching 1938
    • Coveniella M.D.Tindale 1986
    Lomagramma J.Sm. 1841
    • Cheiloepton Fée 1845
    Maxonia C.Chr. 1916 Megalastrum Holttum 1986 Mickelia R.C.Moran, Labiak & Sundue 2010 Olfersia Raddi 1819
    • Dorcapteris C.Presl 1851
    Parapolystichum (Keyserl.) Ching 1940 Phanerophlebia C. Presl 1836 Pleocnemia C.Presl 1836 Polybotrya Humb. & Bonpl. ex Willd. 1810
    • Soromanes Fée 1845
    Polystichopsis (J.Sm.) Holttum 1947 Polystichum Roth 1800
    • Acropelta T.Nakai 1953
    • Adenoderris J.Sm. 1875
    • Aetopteron Ehrh. ex House 1920
    • Hemesteum H.Lév. 1915
    • Hypopeltis Michx. 1803
    • Papuapteris C. Chr. 1937
    • Plecosorus Fée 1852
    • Sorolepidium Christ 1911
    Rumohra Raddi 1819 Stigmatopteris C.Chr. 1909 Teratophyllum Mett. ex Kuhn 1870 Trichoneuron Ching 1965[17]

    Didymochlaena has been removed to Didymochlaenaceae, and Hypodematium and Leucostegia to Hypodematiaceae. Aenigmopteris has at times been suggested to belong to this family, on the grounds of its morphological similarity to Ctenitis, but molecular phylogeny has led to its submersion within Tectaria.[18] Likewise, Dryopolystichum has been placed in Lomariopsidaceae.[19]


    1. ^ a b Christenhusz, Maarten J. M.; Zhang, Xian-Chun; Schneider, Harald (18 February 2011). "A linear sequence of extant families and genera of lycophytes and ferns" (PDF). Phytotaxa. 19: 7–54. ISSN 1179-3163..mw-parser-output cite.citation{font-style:inherit}.mw-parser-output q{quotes:"""""'"'"}.mw-parser-output code.cs1-code{color:inherit;background:inherit;border:inherit;padding:inherit}.mw-parser-output .cs1-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/6/65/Lock-green.svg/9px-Lock-green.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-lock-limited a,.mw-parser-output .cs1-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Lock-gray-alt-2.svg/9px-Lock-gray-alt-2.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/a/aa/Lock-red-alt-2.svg/9px-Lock-red-alt-2.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-subscription,.mw-parser-output .cs1-registration{color:#555}.mw-parser-output .cs1-subscription span,.mw-parser-output .cs1-registration span{border-bottom:1px dotted;cursor:help}.mw-parser-output .cs1-hidden-error{display:none;font-size:100%}.mw-parser-output .cs1-visible-error{font-size:100%}.mw-parser-output .cs1-subscription,.mw-parser-output .cs1-registration,.mw-parser-output .cs1-format{font-size:95%}.mw-parser-output .cs1-kern-left,.mw-parser-output .cs1-kern-wl-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right,.mw-parser-output .cs1-kern-wl-right{padding-right:0.2em}
    2. ^ Sue Olsen. 2007. Encyclopedia of Garden Ferns Timber Press: Portland, OR, USA. ISBN 978-0-88192-819-8
    3. ^ a b c d Smith et al., 2006 Archived February 26, 2008, at the Wayback Machine. Alan R. Smith, Kathleen M. Pryer, Eric Schuettpelz, Petra Korall, Harald Schneider & Paul G. Wolf: "A classification for extant ferns," Taxon, 55(3): 705–731 (Aug 2006)
    4. ^ Eric Schuettpelz and Kathleen M. Pryer. 2009. "Evidence for a Cenozoic radiation of ferns in an angiosperm-dominated canopy". Proceedings of the National Academy of Sciences 106(27):11200-11205. doi:10.1073/pnas.0811136106
    5. ^ Karl U. Kramer (with Richard E. Holttum, Robin C. Moran, and Alan R. Smith). 1990. "Dryopteridaceae". pages ??. In: Klaus Kubitzki (general editor); Karl U. Kramer and Peter S. Green (volume editors) The Families and Genera of Vascular Plants volume I. Springer-Verlag: Berlin;Heidelberg, Germany. ISBN 978-0-387-51794-0
    6. ^ a b Alan R. Smith, Kathleen M. Pryer, Eric Schuettpelz, Petra Korall, Harald Schneider, and Paul G. Wolf. 2008. "Dryopteridaceae". pages ??. In: "Fern Classification". pages 417-467. In: Tom A. Ranker and Christopher H. Haufler (editors). Biology and Evolution of Ferns and Lycophytes. Cambridge University Press. ISBN 978-0-521-87411-3
    7. ^ Hong-Mei Liu, Xian-Chun Zhang, Wei Wang, Yin-Long Qiu, and Zhi-Duan Chen. 2007. "Molecular Phylogeny of the Fern Family Dryopteridaceae inferred from Chloroplast rbcL and atpB genes". International Journal of Plant Sciences 168(9):1311-1323. doi:10.1086/521710
    8. ^ Robbin C. Moran, Paulo H. Labiak, and Michael Sundue. 2010. "Phylogeny and character evolution of the bolbitidoid ferns (Dryopteridaceae)". International Journal of Plant Sciences 171(5):547-559. doi:10.1086/652191
    9. ^ Robbin C. Moran, Paulo H. Labiak, and Michael Sundue. 2010. "Synopsis of Mickelia, a newly recognized genus of bolbitidoid ferns (Dryopteridaceae)". Brittonia 62(4):337-356.
    10. ^ a b c d e f g Li-Bing Zhang, Liang Zhang, Shi-Yong Dong, and Atsushi Ebihara. 2012. "Molecular circumscription and major evolutionary lineages of the fern genus Dryopteris (Dryopteridaceae)". BMC Evolutionary Biology 12(1):180
    11. ^ a b c d e He H, Wu SG, Xiang JY, Barrington DS (2013) "Arachniodes". In: Wu ZY, Raven PH, Hong DY (eds) Flora of China, vol 2–3.
    12. ^ Christenhusz & Chase 2014.
    13. ^ Pteridophyte Phylogeny Group (November 2016). "A community-derived classification for extant lycophytes and ferns". Journal of Systematics and Evolution. 54 (6): 563–603. doi:10.1111/jse.12229.
    14. ^ Family:Dryopteridaceae USDA-ARS Germplasm Resources Information Network (GRIN) 25 Jan 2012
    15. ^ Dryopteridaceae Tropicos.org. Missouri Botanical Garden. 25 Jan 2012
    16. ^ "Nothoperanema". Natural Resources Conservation Service PLANTS Database. USDA. Retrieved 25 January 2012.
    17. ^ Hong-Mei Liu, Xian-Chun Zhang, Mei-Ping Wang, Hui Shang, Shi-Liang Zhou, Yue-Hong Yan, Xue-Ping Wei, Wen-Bin Xu, Harald Schneider. 2016. "Phylogenetic placement of the enigmatic fern genus Trichoneuron informs on the infra-familial relationship of Dryopteridaceae". Plant Systematics and Evolution 302:319–332
    18. ^ Chen, Cheng-Wei; RothfelsE, Carl J.; Mustapeng, Andi Maryani A.; Gubilil, Markus; Karger, Dirk Nikolaus; Kessler, Michael; Huang, Yao-Moan. "End of an enigma: Aenigmopteris belongs in Tectaria (Tectariaceae: Polypodiopsida)". Journal of Plant Research. doi:10.1007/s10265-017-0966-9.
    19. ^ Chen, Cheng-Wei; Sundue, Michael; Kuo, Li-Yaung; Teng, Wei-Chih; Huang, Yao-Moan. "Phylogenetic analyses place the monotypic Dryopolystichum within Lomariopsidaceae". PhytoKeys. 78: 83–107. doi:10.3897/phytokeys.78.12040.