View source for Dryopteridaceae

{{Short description|Family of ferns}}
{{Automatic taxobox
| image = Dryopteris carthusiana.jpg
| image_caption = ''[[Dryopteris carthusiana]]''
| taxon = Dryopteridaceae
| authority = [[Wilhelm Gustav Franz Herter|Herter]] (''nom. cons.'')
| subdivision_ranks = Subfamilies
| subdivision = 
* Dryopteridoideae
* [[Elaphoglossoideae]]
* [[Polybotryoideae]]
| synonyms =
* Bolbitidaceae <small>Ching 1978</small>
* Elaphoglossaceae <small>Pichi-Sermolli 1968</small>
* Filicaceae <small>de Jussieu 1789 nom. ill.</small>
* Peranemataceae <small>Ching 1940 non Buetschli 1884</small>
}}

The '''Dryopteridaceae''' are a [[Family (biology)|family]] of [[leptosporangiate fern]]s in the [[Order (biology)|order]] [[Polypodiales]]. They are known [[Common name|colloquially]] as the '''wood ferns'''. In the [[Pteridophyte Phylogeny Group]] classification of 2016 (PPG&nbsp;I), the family is placed in the suborder [[Polypodiineae]].<ref name=PPGI/> Alternatively, it may be treated as the subfamily '''Dryopteridoideae ''' of a very broadly defined family [[Polypodiaceae]] ''[[Sensu|sensu lato]]''.<ref name=ChriChas14/>

The family contains about 1700 [[species]] and has a [[cosmopolitan distribution]]. Species may be [[terrestrial plant|terrestrial]], [[epipetric]], [[hemiepiphyte|hemiepiphytic]], or [[epiphytic]]. Many are [[Gardening|cultivated]] as [[ornamental plant]]s.<ref name=Olsen-2007/>  The largest [[genera]] are ''Elaphoglossum'' (600+), ''Polystichum'' (260), ''Dryopteris'' (225), and ''Ctenitis'' (150). These four genera contain about 70% of the species.<ref name=Smith-2006/>  Dryopteridaceae [[Evolutionary radiation|diverged]] from the other families in [[eupolypods I]] about 100 million years ago.<ref name=Schuettpelz-2009/>
The fossil record appears to show the earliest members of the group emerging some 99 MYA during the Mid Cretaceous.<ref>{{Cite journal |last1=Long |first1=Xiaoxuan |last2=Peng |first2=Yuan |last3=Feng |first3=Qi |last4=Engel |first4=Michael S. |last5=Shi |first5=Chao |last6=Wang |first6=Shuo |date=2023-09-01 |title=A new fossil fern of the Dryopteridaceae (Polypodiales) from the mid-Cretaceous Kachin amber |url=https://link.springer.com/article/10.1007/s12549-023-00572-4 |journal=Palaeobiodiversity and Palaeoenvironments |language=en |volume=103 |issue=3 |pages=489–494 |doi=10.1007/s12549-023-00572-4 |bibcode=2023PdPe..103..489L |issn=1867-1608|url-access=subscription }}</ref> 

== Description ==
The [[rhizomes]] are often stout, creeping, ascending, or erect, and sometimes [[scandent]] or [[Vine|climbing]], with non[[clathrate]] [[Trichome|scales]] at [[Glossary of botanical terms#A|apices]]. [[Frond]]s are usually monomorphic, less often [[Frond dimorphism|dimorphic]], or sometimes scaly or [[Gland (botany)|glandular]], but less commonly hairy. [[Petiole (botany)|Petioles]] have numerous round, [[Vascular tissue|vascular bundles]] arranged in a ring, or rarely as few as three; the [[Glossary of botanical terms#A|adaxial]] bundles are largest. [[Leaf#Veins|Veins]] are [[pinnate]] or [[Divaricate|forking]], free to variously [[Anastomosis|anastomosing]]; the [[areole]]s occur with or without included veinlets; [[Sorus|sori]] are usually round, [[Glossary of botanical terms#A|acrostichoid]] (covering the entire abaxial surface of the lamina) in a few [[Clade|lineages]]; usually [[indusium|indusiate]], or sometimes exindusiate. Indusia, when present, are round-[[Reniform leaf|reniform]] or [[peltate]]. [[Sporangia]] have three-rowed, short to long stalks; [[spore]]s are reniform, [[Spore#Anatomy|monolete]], [[spore perine|perine]] or [[Glossary of botanical terms#W|winged]].<ref name=Smith-2006/>

==Taxonomy==

=== History ===
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]].<ref name=Kramer-1990/> [[Molecular phylogenetic]] studies found Kramer's [[Circumscription (taxonomy)|version]] of the Dryopteridaceae to be [[polyphyletic]], and it was [[Segregate (taxonomy)|split up]] by Smith and others in 2006.<ref name=Smith-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.<ref name=Smith-2008/>  Some authors have already treated these genera as outside of the Dryopteridaceae.<ref name=Christenhusz-2011/>

In 2007, a [[phylogenetic]] study of [[DNA sequence]]s showed that ''Pleocnemia'' should be transferred from the Tectariaceae to the Dryopteridaceae.<ref name=Liu-2007/> In 2010, in a paper on [[bolbitidoid]] ferns, ''Arthrobotrya'' was resurrected from ''Teratophyllum''.<ref name=Moran-2010a/> Later that year, ''Mickelia'' was described as a new genus.<ref name=Moran-2010b/>

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 [[Genus#Generic name|generic name]].<ref name=Smith-2008/>

In 2012, a phylogenetic study of ''Dryopteris'' and its relatives included ''Acrophorus'', ''Acrorumohra'', ''Diacalpe'', ''Dryopsis'', ''Nothoperanema'', and ''Peranema'' within that genus.<ref name=Zhang-2012/> The ''Flora of China'' treatment of the family, published in 2013, used phylogenetic results to sink ''Lithostegia'' and ''Phanerophlebiopsis'' into ''Arachniodes''.<ref name=FoC-Arachniodes />

The Dryopteridaceae <small>[[Wilhelm Gustav Franz Herter|Herter]]</small>, under the classification system of Christenhusz and Chase (2014), were submerged as subfamily Dryopteridoideae <small>[[Johann Heinrich Friedrich Link|Link]]</small>, 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.<ref name=PPGI/>

===Phylogeny===
The following cladogram for the suborder [[Polypodiineae]] (eupolypods I), based on the consensus cladogram in the [[Pteridophyte Phylogeny Group]] classification of 2016 (PPG I),<ref name=PPGI/> shows a likely [[phylogenetics|phylogenetic]] relationship between Dryopteridaceae and the other families of the clade.
{{Polypodiineae cladogram}}

=== Subdivision ===
{{cladogram|title=Phylogeny of Dryopteridaceae<ref>{{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= |article-number=909768 | doi=10.3389/fpls.2022.909768 |pmid= 36092417|pmc=9449725 |bibcode= 2022FrPS...1309768N|doi-access=free}}</ref><ref>{{cite web |last1= |first1= |display-authors=et al. |year=2024 |title=Tree viewer: interactive visualization of FTOL |url=https://fernphy.github.io/viewer.html |version=FTOL v1.7.0 ([[GenBank]] release 261) |access-date=1 May 2025}}</ref>|
{{clade|style=font-size:90%;line-height:80%;width:400px
|1={{Clade
  |1={{Clade
    |label1=Elaphoglossoideae
    |1={{Clade
      |label1=Lastreopsideae
      |1={{Clade
        |1=''[[Parapolystichum]]''
        |2={{Clade
          |1=''[[Lastreopsis]]''
          |2={{Clade
            |1=''[[Rumohra]]''
            |2=''[[Megalastrum]]''
             }}
           }}
         }}
      |label2=Bolbitideae
      |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 (plant)|Olfersia]]''
          |2={{Clade
            |1=''[[Maxonia]]''
            |2={{Clade
              |1=''[[Cyclodium]]''
              |2=''[[Polybotrya]]''
               }}
             }}
           }}
         }}
       }}
    |label2=Dryopteridoideae
    |2={{Clade
      |1={{Clade
      |label1=Ctenitideae
      |1=''[[Ctenitis]]''
         }}
      |2={{Clade
        |label1=Polysticheae
        |1={{Clade
          |1={{Clade
            |1=''[[Phanerophlebia (plant)|Phanerophlebia]]''
            |2=''[[Cyrtomium]]''
             }}
          |2=''[[Polystichum]]''
           }}
        |label2=Dryopterideae
        |2={{Clade
          |1=''[[Arachniodes]]''
          |2=''[[Dryopteris]]''
           }}
         }}
       }}
     }}
   }}
}}
}}

The PPG I classification divides the family into three subfamilies, listed below.<ref name=PPGI>{{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|bibcode=2016JSyEv..54..563. |s2cid=39980610|doi-access=free}}</ref>
* Subfamily [[Polybotryoideae]] <small>H.M.Liu & X.C.Zhang</small>
** ''[[Cyclodium]]'' <small>C.Presl</small>
** ''[[Maxonia]]'' <small>C.Chr.</small>
** ''[[Olfersia (plant)|Olfersia]]'' <small>Raddi</small>
** ''[[Polybotrya]]'' <small>Humb. & Bonpl. ex Willd.</small>
** ''[[Polystichopsis]]'' <small>(J.Sm.) Holttum</small>
** ''[[Stigmatopteris]]'' <small>C.Chr.</small>
** ''[[Trichoneuron]]'' <small>Ching</small>
* Subfamily [[Elaphoglossoideae]] <small>(Pic.Serm.) Crabbe, Jermy & Mickel</small>
** ''[[Arthrobotrya]]'' <small>J.Sm.</small>
** ''[[Bolbitis]]'' <small>Schott</small>
** ''[[Elaphoglossum]]'' <small>Schott ex J.Sm.</small>
** ''[[Lastreopsis]]'' <small>Ching</small>
** ''[[Lomagramma]]'' <small>J.Sm.</small>
** ''[[Megalastrum]]'' <small>Holttum</small>
** ''[[Mickelia]]'' <small>R.C.Moran, Labiak & Sundue</small>
** ''[[Parapolystichum]]'' <small>(Keyserl.) Ching</small>
** ''[[Pleocnemia]]'' <small>C.Presl</small>
** ''[[Rumohra]]'' <small>Raddi</small>
** ''[[Teratophyllum]]'' <small>Mett. ex Kuhn</small>
* Subfamily Dryopteridoideae <small>Link</small>
** ''[[Arachniodes]]'' <small>Blume</small>
** ''[[Ctenitis]]''  <small>(C.Chr.) C.Chr.</small>
** ''[[Cyrtomium]]'' <small>C.Presl</small>
** ''[[Dryopteris]]'' <small>Adans.</small>
** ''[[Phanerophlebia (plant)|Phanerophlebia]]'' <small>C.Presl</small>
** ''[[Polystichum]]'' <small>Roth</small>

''[[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).<ref>{{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 |bibcode=2018JPlR..131...67C |s2cid=4573970 }}</ref> ''[[Dryopolystichum]]'' has been placed in [[Lomariopsidaceae]].<ref>{{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 |bibcode=2017PhytK..78...83C |doi-access=free }}</ref>

==References==
{{Reflist|30em|refs=
<ref name=ChriChas14>{{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}}</ref>

<ref name=Kramer-1990>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}}</ref>

<ref name=Smith-2006>[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)</ref>

<ref name=Liu-2007>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}}</ref>

<ref name=Olsen-2007>Sue Olsen. 2007. ''Encyclopedia of Garden Ferns'' Timber Press: Portland, OR, USA. {{ISBN|978-0-88192-819-8}}</ref>

<ref name=Smith-2008>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}}</ref>

<ref name=Schuettpelz-2009>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}}</ref>

<ref name=Moran-2010a>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}}</ref>

<ref name=Moran-2010b>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.</ref>

<ref name=Christenhusz-2011>{{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 | bibcode=2011Phytx..19....7C | hdl=10138/28042 }}</ref>

<ref name=Zhang-2012>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</ref>

<ref name=FoC-Arachniodes>He H, Wu SG, Xiang JY, Barrington DS (2013) "Arachniodes". In: Wu ZY, Raven PH, Hong DY (eds) ''Flora of China'', vol 2–3.</ref>
}}

== Bibliography ==
{{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 (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|bibcode=2006Taxon..55..705S |ref={{harvid|Smith et al|2006}} }}
{{Refend}}

== External links ==
* [http://www.mapress.com/phytotaxa/ ''Phytotaxa'']

{{Plant classification}}
{{Fern classification}}
{{Taxonbar|from=Q656683}}
{{Authority control}}

[[Category:Dryopteridaceae| ]]
[[Category:Fern families]]