Dryopteridaceae 

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 are reniform, monolete, perine or winged.

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. Molecular phylogenetic studies found Kramer's version of the Dryopteridaceae to be polyphyletic, and it was split up by Smith and others in 2006. 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. Some authors have already treated these genera as outside of the Dryopteridaceae.

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

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.

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

The Dryopteridaceae Herter, under the classification system of Christenhusz and Chase (2014), were submerged as subfamily Dryopteridoideae Link, one of eight subfamilies constituting family Polypodiaceae. This family corresponds to the clade eupolypods I.{{sfn|Christenhusz|Chase|2014}} The Pteridophyte Phylogeny Group classification of 2016 (PPG I) retained the family.

The following cladogram for the suborder Polypodiineae (eupolypods I), based on the consensus cladogram in the Pteridophyte Phylogeny Group classification of 2016 (PPG I), shows a likely phylogenetic relationship between Dryopteridaceae and the other families of the clade.

{{Polypodiineae cladogram}}

{{cladogram|title=Phylogeny of Dryopteridaceae{{cite journal |last1=Nitta |first1=Joel H. |last2=Schuettpelz |first2=Eric |last3=Ramírez-Barahona |first3=Santiago |last4=Iwasaki |first4=Wataru |display-authors=et al. |year=2022 |title=An Open and Continuously Updated Fern Tree of Life |journal=Frontiers in Plant Science |volume=13 |issue= |page= | doi=10.3389/fpls.2022.909768 |pmid= 36092417|pmc=9449725 |bibcode= |doi-access=free}}{{cite web|last1= |first1= |last2= |first2= |last3= |first3= |last4= |first4= |display-authors=et al. |year=2022 |title=Tree viewer: interactive visualization of FTOL |url=https://fernphy.github.io/viewer.html |version=FTOL v1.3.0 |access-date=12 December 2022}}|

{{clade|style=font-size:90%;line-height:80%;width:300px

|1={{Clade

|1={{Clade

|label1=Elaphoglossoideae

|1={{Clade

|1={{Clade

|1=Parapolystichum

|2={{Clade

|1=Lastreopsis

|2={{Clade

|1=Rumohra

|2=Megalastrum

}}

}}

}}

|2={{Clade

|1=Pleocnemia

|2={{Clade

|1=Bolbitis

|2={{Clade

|1={{Clade

|1=Lomagramma

|2={{Clade

|1=Arthrobotrya

|2=Teratophyllum

}}

}}

|2={{Clade

|1=Mickelia

|2=Elaphoglossum

}}

}}

}}

}}

}}

}}

|2={{Clade

|label1=Polybotryoideae

|1={{Clade

|1=Stigmatopteris

|2={{Clade

|1={{Clade

|1=Trichoneuron

|2=Polystichopsis

}}

|2={{Clade

|1=Olfersia

|2={{Clade

|1=Maxonia

|2={{Clade

|1=Cyclodium

|2=Polybotrya

}}

}}

}}

}}

}}

|label2=Dryopteridoideae

|2={{Clade

|1=Ctenitis

|2={{Clade

|1={{Clade

|1={{Clade

|1=Phanerophlebia

|2=Cyrtomium

}}

|2=Polystichum

}}

|2={{Clade

|1=Arachniodes

|2=Dryopteris

}}

}}

}}

}}

}}

}}

}}

The PPG I classification divides the family into three subfamilies, listed below.{{cite journal|last1=Pteridophyte Phylogeny Group|title=A community-derived classification for extant lycophytes and ferns|journal=Journal of Systematics and Evolution|date=November 2016|volume=54|issue=6|pages=563–603|doi=10.1111/jse.12229|s2cid=39980610|doi-access=free}}

• Subfamily Polybotryoideae H.M.Liu & X.C.Zhang
• Cyclodium C.Presl
• Maxonia C.Chr.
• Olfersia Raddi
• Polybotrya Humb. & Bonpl. ex Willd.
• Polystichopsis (J.Sm.) Holttum
• Stigmatopteris C.Chr.
• Trichoneuron Ching
• Subfamily Elaphoglossoideae (Pic.Serm.) Crabbe, Jermy & Mickel
• Arthrobotrya J.Sm.
• Bolbitis Schott
• Elaphoglossum Schott ex J.Sm.
• Lastreopsis Ching
• Lomagramma J.Sm.
• Megalastrum Holttum
• Mickelia R.C.Moran, Labiak & Sundue
• Parapolystichum (Keyserl.) Ching
• Pleocnemia C.Presl
• Rumohra Raddi
• Teratophyllum Mett. ex Kuhn
• Subfamily Dryopteridoideae Link
• Arachniodes Blume
• Ctenitis (C.Chr.) C.Chr.
• Cyrtomium C.Presl
• Dryopteris Adans.
• Phanerophlebia C.Presl
• Polystichum Roth

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 (Tectariaceae).{{cite journal | first1=Cheng-Wei |last1=Chen |first2=Carl J. |last2=RothfelsE |first3=Andi Maryani A. |last3=Mustapeng |first4=Markus |last4=Gubilil |first5=Dirk Nikolaus |last5=Karger |first6=Michael |last6=Kessler |first7=Yao-Moan |last7=Huang |title=End of an enigma: Aenigmopteris belongs in Tectaria (Tectariaceae: Polypodiopsida) |journal=Journal of Plant Research |volume=131 |pages=67–76 |doi=10.1007/s10265-017-0966-9|year=2018 |issue=1 |pmid=28741041 |s2cid=4573970 }} Dryopolystichum has been placed in Lomariopsidaceae.{{cite journal |first1=Cheng-Wei |last1=Chen |first2=Michael |last2=Sundue |first3=Li-Yaung |last3=Kuo |first4=Wei-Chih |last4=Teng |first5=Yao-Moan |last5=Huang |title=Phylogenetic analyses place the monotypic Dryopolystichum within Lomariopsidaceae | journal=PhytoKeys |issue=78 |pages=83–107 |doi=10.3897/phytokeys.78.12040|year=2017 |pmid=28781553 |pmc=5543276 |doi-access=free }}

{{Reflist|30em|refs=

{{Cite journal |last1=Christenhusz |first1=Maarten J.M. |last2=Chase |first2=Mark W. |date=2014 |title=Trends and concepts in fern classification |journal=Annals of Botany |volume=113 |issue=9 |pages=571–594 |doi=10.1093/aob/mct299 |pmid=24532607 |pmc=3936591 |name-list-style=amp}}

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}}

[http://www.pryerlab.net/publication/fichier749.pdf Smith et al., 2006] {{webarchive |url=https://web.archive.org/web/20080226232147/http://www.pryerlab.net/publication/fichier749.pdf |date=February 26, 2008 }} 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)

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}}

Sue Olsen. 2007. Encyclopedia of Garden Ferns Timber Press: Portland, OR, USA. {{ISBN|978-0-88192-819-8}}

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}}

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}}

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}}

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.

{{cite journal | first1=Maarten J. M. | last1=Christenhusz | first2=Xian-Chun | last2=Zhang | first3=Harald | last3=Schneider | title=A linear sequence of extant families and genera of lycophytes and ferns | journal=Phytotaxa | volume=19 | pages=7–54 | date=18 February 2011 | url=http://www.mapress.com/phytotaxa/content/2011/f/pt00019p054.pdf | issn=1179-3163| doi=10.11646/phytotaxa.19.1.2 | hdl=10138/28042 }}

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

He H, Wu SG, Xiang JY, Barrington DS (2013) "Arachniodes". In: Wu ZY, Raven PH, Hong DY (eds) Flora of China, vol 2–3.

}}

{{Refbegin|30em}}

• {{cite journal|last1=Smith|first1=Alan R.|last2=Pryer|first2=Kathleen M.|last3=Schuettpelz|first3=Eric|last4=Korall|first4=Petra|last5=Schneider|first5=Harald|last6=Wolf|first6=Paul G.|title=A Classification for Extant Ferns|journal=Taxon|date=1 January 2006|volume=55|issue=3|pages=705–731|url=https://www.idigbio.org/wiki/images/9/95/Smith_et_al_2006.pdf|jstor=25065646|doi=10.2307/25065646|ref={{harvid|Smith et al|2006}} }}

{{Refend}}

• [http://www.mapress.com/phytotaxa/ Phytotaxa]

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Category:Fern families