View source for Blechnaceae

{{Short description|Family of ferns}}
{{Automatic taxobox
| image = Blechnum spicant2.jpg
| image_caption = ''[[Struthiopteris spicant]]''
| taxon = Blechnaceae
| authority = Newman{{sfn|PPG I|2016}}
| subdivision_ranks = Subfamilies and genera
| subdivision = [[#Subfamilies and genera|See text]]
| type_genus = ''[[Blechnum]]''
| synonyms =
* Stenochlaenaceae <small>Ching 1978</small>
| synonyms_ref = {{sfn|Christenhusz|Chase|2014}}
}}
{{Commons category|Blechnaceae}}
[[File:擬烏毛蕨 - 溪頭自然教育園區 Blechnopsis orientalis - Xitou Nature Education Area, Taiwan 20220628191131 08.jpg|thumb|x300px|''[[Blechnopsis orientalis]]''. The young opening fronds of many species are usually tinged with red. ]]

'''Blechnaceae''' is a family of [[fern]]s in the order [[Polypodiales]], with a [[cosmopolitan distribution]]. Its status as a family and the number of genera included have both varied considerably. In the [[Pteridophyte Phylogeny Group]] classification of 2016 (PPG I), the family has 24 genera, and excludes genera placed in the separate family [[Onocleaceae]]. The family is divided into three subfamilies, including Blechnoideae [[Sensu|s.s.]]{{sfn|PPG I|2016}} Alternatively, the entire family may be treated as the subfamily Blechnoideae [[Sensu|s.l.]] of a very broadly defined family [[Aspleniaceae]], and include genera others place in Onocleaceae.{{sfn|Christenhusz|Chase|2014}}

== Description ==
Most are ground dwelling, some are climbers, such as ''[[Stenochlaena]]''. A characteristic feature of many species is that the young opening [[frond]]s are usually tinged with red.

== Taxonomy ==
The family was created by [[Edward Newman (entomologist)|Newman]] in 1844. In 2014, Christenhusz and Chase submerged it as subfamily Blechnoideae within the family [[Aspleniaceae]] and included [[Onocleaceae]] in it.{{sfn|Christenhusz|Chase|2014}} The [[Pteridophyte Phylogeny Group#PPG I|PPG I]] classification of 2016 restored it to family status.{{sfn|PPG I|2016}}

Blechnaceae is a member of the eupolypods II clade (now the suborder Aspleniineae), in the order [[Polypodiales]].<ref name=Rothfels-2012/> It is related to other families in the clade as in the following cladogram:<ref name=Lehtonen-2011/><ref name=Rothfels-2012/>
{{eupolypods II cladogram}}

=== Subdivisions ===
The number of genera accepted within Blechnaceae (or Blechnoideae when treated as a subfamily) has varied between authors. Christenhusz and Chase (2014), treating the group as a subfamily, describe the situation as follows: "Blechnoideae comprise three major clades, one corresponding to ''[[Onoclea]]'' ''sensu lato'', a second corresponding to ''[[Woodwardia]]'', sister to all other species that can be treated as the single genus ''[[Blechnum]]''. However, the subclade sister to the rest of ''Blechnum'' ''sensu lato'' contains the vining taxa ''Stenochlaena'', ''[[Salpichlaena]]'' <small>J.Sm.</small> and a few non-vining ''Blechnum'' species with long-creeping rhizomes, which may have to be accepted at the generic level pending further studies. ''[[Brainea]]'', ''[[Doodia]]'', ''[[Pteridoblechnum]]'' and ''[[Sadleria]]'' belong to ''Blechnum'' sensu lato."{{sfn|Christenhusz|Chase|2014}}

Perrie et al. (2014) retained the family rank and excluded ''[[Onoclea]]'' as a separate family, [[Onocleaceae]], sister to Blechnaceae. They identified three major clades within the remaining Blechnaceae, which they labelled ''Woodwardia'', super-''Stenochlaena'' and super-''Blechnum'', with the latter two as [[sister groups]]. They did not consider ''[[Blechnum]]'' as monophyletic and recommended revision of intergeneric boundaries, resulting in seven genera.{{sfn|Perrie et al.|2014}}

Gasper et al. (2016), independently of Christenhusz and Chase, examined the deeper relationships of the 
Blechnaceae, retaining its family status and excluding Onocleaceae, and allocated the three major clades of Perrie et al. (2014) to subfamilies: Woodwardioideae, Stenochlaenoideae and Blechnoideae. Their approach to the [[polyphyletic]] nature of ''Blechnum'' was to create a series of monophyletic [[segregate (taxonomy)|segregate]] genera, resulting in 24 genera in total.{{sfn|Gasper et al.|2016a}}{{sfn|Gasper et al.|2016b}} The relationship between the three subfamilies was found to be as follows:{{sfn|Gasper et al.|2016a}}
{{clade
|label1=Blechnaceae
|1={{clade
   |1=Woodwardioideae
   |2={{clade
      |1=Stenochlaenoideae
      |2=Blechnoideae
      }}
   }}
}}

The approximate relationship between some of the taxa used in the classifications of Christenhusz and Chase (2014){{sfn|Christenhusz|Chase|2014}} and PPG I (2016){{sfn|PPG I|2016}} is shown in the table below.
{| class="wikitable"
|+ Approximate relationships in two classifications
|-
! Christenhusz & Chase (2014) !! PPG I (2016)
|-
| Family Aspleniaceae
:Subfamily Blechnoideae
::Genus ''Blechnum'' s.l.
:::&nbsp;
| Suborder [[Aspleniineae]]
:Family Blechnaceae
::Subfamily Blechnoideae
:::(containing ''[[Blechnum]]'' s.s.)
|}

===Subfamilies and genera===
{{cladogram|title=Phylogeny of Blechnaceae<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= |page= 909768| doi=10.3389/fpls.2022.909768 |pmid= 36092417|pmc= 9449725|bibcode= |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:450px
|1={{Clade
  |1={{Clade
    |label1=Woodwardioideae
    |1={{Clade
      |1=''[[Lorinseria]]''
      |2={{Clade
        |1=''[[Anchistea]]''
        |2=''[[Woodwardia]]''
         }}
       }}
     }}
  |2={{Clade
    |label1=Stenochlaenoideae
    |1={{Clade
      |1=''[[Salpichlaena]]''
      |2={{Clade
        |1=''[[Stenochlaena]]''
        |2=''[[Telmatoblechnum]]''
         }}
       }}
    |label2=Blechnoideae
    |2={{Clade
      |1=''[[Brainea]]''
      |2={{Clade
        |1={{Clade
        |label1=Struthiopterideae
        |1={{Clade
          |1=''[[Spicantopsis]]''
          |2={{Clade
            |1=''[[Blechnidium]]''
            |2=''[[Struthiopteris]]''
             }}
           }}
           }}
        |2={{Clade
          |1={{Clade
          |label1=Sadlerieae
          |1={{Clade
            |1={{Clade
              |1=''[[Cleistoblechnum]]''
              |2=''[[Blechnopsis]]''
               }}
            |2=''[[Sadleria]]''
             }}
             }}
          |2={{Clade
            |1=''[[Lomaridium]]''
            |2={{Clade
              |1={{Clade
                |label1=Lomarieae
                |1=''[[Lomaria]]''
                 }}
              |2={{Clade
                |1=''[[Lomariocycas]]''
                |2={{Clade
                  |label1=Blechneae
                  |1={{Clade
                    |1=''[[Cranfillia]]''
                    |2={{Clade
                      |1=''[[Icarus (plant)|Icarus]]''
                      |2={{Clade
                        |1=''[[Blechnum]]''
                        |2=''[[Austroblechnum]]''
                         }}
                       }}
                     }}
                  |label2=Doodieae
                  |2={{Clade
                    |1={{Clade
                      |1=''[[Diploblechnum]]''
                      |2={{Clade
                        |1={{Clade
                          |1=''[[Neoblechnum]]''
                          |2={{Clade
                            |1=''[[Doodia maxima]]''
                            |2=''[[Oceaniopteris]]''
                             }}
                           }}
                        |2={{Clade
                          |1=''Oceaniopteris'' species-group 2
                          |2=''[[Doodia]]''
                           }}
                         }}
                       }}
                    |2=''[[Parablechnum]]''
                     }}
                   }}
                 }}
               }}
             }}
           }}
         }}
       }}
     }}
   }}
}}
}}

In 2016, the Pteridophyte Phylogeny Group followed Gasper et al. (2016) in accepting 24 genera, grouped into three subfamilies with approximately 265 species, most of which are placed in the subfamily Blechnoideae.{{sfn|PPG I|2016}}

Subfamily Woodwardioideae <small>Gasper, V.A.O.Dittrich & Salino</small>
*''[[Anchistea]]'' <small>C.Presl</small>
*''[[Lorinseria]]'' <small>C.Presl</small>
*''[[Woodwardia]]'' <small>Sm.</small>
Subfamily Stenochlaenoideae <small>(Ching) J.P.Roux</small>
*''[[Salpichlaena]]'' <small>J.Sm.</small>
*''[[Stenochlaena]]'' <small>J.Sm.</small>
*''[[Telmatoblechnum]]'' <small>Perrie, D.J.Ohlsen & Brownsey</small>
Subfamily Blechnoideae <small>Gasper, V.A.O.Dittrich & Salino</small>
*''[[Austroblechnum]]'' <small>Gasper & V.A.O.Dittrich</small>
*''[[Blechnidium]]'' <small>T.Moore</small>
*''[[Blechnopsis]]'' <small>C.Presl</small>
*''[[Blechnum]]'' <small>L.</small>
*''[[Brainea]]'' <small>J.Sm.</small>
*''[[Cleistoblechnum]]'' <small>Gasper & Salino</small>
*''[[Cranfillia]]'' <small>Gasper & V.A.O.Dittrich</small>
*''[[Diploblechnum]]'' <small>Hayata</small>
*''[[Doodia]]'' <small>R.Br.</small>
*''[[Icarus (plant)|Icarus]]'' <small>Gasper & Salino</small>
*''[[Lomaria]]'' <small>Willd.</small>
*''[[Lomaridium]]'' <small>C.Presl</small>
*''[[Lomariocycas]]'' <small>(J.Sm.) Gasper & A.R.Sm.</small>
*''[[Neoblechnum]]'' <small>Gasper & V.A.O.Dittrich</small>
*×''[[Oceanidoodia]]'' <small>Liu, Schuettp. & H. Schneid.</small> 
*''[[Oceaniopteris]]'' <small>Gasper & Salino</small>
*''[[Parablechnum]]'' <small>C.Presl</small>
*''[[Sadleria]]'' <small>Kaulf.</small>
*''[[Struthiopteris]]'' <small>Scop.</small>
Subfamily ''incertae sedis''
*{{extinct}}''[[Trawetsia]]'' {{small|Smith ''et al''}} [[Ypresian]], [[Princeton Chert]]<ref name="Trawetsia2006">{{cite journal |last1=Smith |first1=S. Y. |last2=Stockey |first2=R. A. |last3=Nishida |first3=H. |last4=Rothwell |first4=G. W. |year=2006 |title=''Trawetsia princetonensis'' gen. et sp. nov.(Blechnaceae): a permineralized fern from the Middle Eocene Princeton Chert |journal=International Journal of Plant Sciences |volume=167 |issue=3 |pages=711–719|doi=10.1086/501034 |s2cid=85160532}}</ref>

==References==
{{Reflist|30em|refs=

<ref name=Lehtonen-2011>{{cite journal
|author=Samuli Lehtonen
|year=2011
|title=Towards Resolving the Complete Fern Tree of Life
|journal=PLOS ONE
|volume=6
|issue=10
|pages=e24851
|doi=10.1371/journal.pone.0024851
|pmid=22022365
|pmc=3192703
|bibcode=2011PLoSO...624851L
|editor1-last=Steinke
|editor1-first=Dirk
|doi-access=free
}}</ref>

<ref name=Rothfels-2012>{{cite journal
|author1=Carl J. Rothfels |author2=Anders Larsson |author3=Li-Yaung Kuo |author4=Petra Korall |author5=Wen- Liang Chiou |author6=Kathleen M. Pryer |year=2012
|title=Overcoming Deep Roots, Fast Rates, and Short Internodes to Resolve the Ancient Rapid Radiation of Eupolypod II Ferns
|journal=Systematic Biology |volume=61 |issue=1 |pages=490–509
|pmid=22223449
|doi=10.1093/sysbio/sys001|doi-access=free }}</ref>
}}

== Bibliography ==
{{refbegin|30em}}
*{{cite journal
|author=Maarten J. M. Christenhusz
|author2=Xian-Chun Zhang
|author3=Harald Schneider
|name-list-style=amp
|year=2011
|title=A linear sequence of extant families and genera of lycophytes and ferns
|journal=[[Phytotaxa]]|volume=19|pages=7–54
|doi=10.11646/phytotaxa.19.1.2
|url=http://www.mapress.com/phytotaxa/content/2011/f/pt00019p054.pdf}}
* {{Cite journal|last1=Christenhusz |first1=Maarten J.M.|authorlink1=Maarten J. M. Christenhusz  |last2=Chase|first2=Mark W.|authorlink2=Mark W. Chase |year=2014|title=Trends and concepts in fern classification|journal=[[Annals of Botany]]|volume=113
|issue=9|pages=571–594|name-list-style=amp|doi=10.1093/aob/mct299  |pmid=24532607  |pmc=3936591}}
* {{cite journal|last1=Gasper|first1=André L. de|last2=Almeida|first2=Thaís E.|last3=Dittrich|first3=Vinícius A. de O.|last4=Smith|first4=Alan R.|last5=Salino|first5=Alexandre|title=Molecular phylogeny of the fern family Blechnaceae (Polypodiales) with a revised genus-level treatment|url=https://www.researchgate.net/publication/309285163|journal=[[Cladistics (journal)|Cladistics]]|date=October 2016|volume=33|issue=4|pages=429–446|doi=10.1111/cla.12173|pmid=34715732 |s2cid=88680078|ref={{harvid|Gasper et al.|2016a}}|doi-access=free}}
* {{cite journal|last1=Gasper|first1=André Luís de|last2=Dittrich|first2=Vinícius Antonio de Oliveira|last3=Smith|first3=Alan Reid|last4=Salino|first4=Alexandre|title=A classification for Blechnaceae (Polypodiales: Polypodiopsida): New genera, resurrected names, and combinations|journal=[[Phytotaxa]]|date=21 September 2016|volume=275|issue=3|pages=191|doi=10.11646/phytotaxa.275.3.1|url=https://www.researchgate.net/publication/308389609|ref={{harvid|Gasper et al.|2016b}}}}
* {{cite journal|last1=Perrie|first1=Leon R.|last2=Wilson|first2=Ruby K.|last3=Shepherd|first3=Lara D.|last4=Ohlsen|first4=Daniel J.|last5=Batty|first5=Erin L.|last6=Brownsey|first6=Patrick J.|last7=Bayly|first7=Michael J.|title=Molecular phylogenetics and generic taxonomy of Blechnaceae ferns|journal=[[Taxon (journal)|Taxon]]|date=28 August 2014|volume=63|issue=4|pages=745–758|doi=10.12705/634.13|ref={{harvid|Perrie et al.|2014}}}}
* {{cite journal |author=Pteridophyte Phylogeny Group I |year=2016 |title=A community-derived classification for extant lycophytes and ferns |journal=Journal of Systematics and Evolution |volume=54 |issue=6 |pages=563–603 |doi=10.1111/jse.12229|s2cid=39980610 |ref={{harvid|PPG I|2016}}|doi-access=free }}
* {{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|ref={{harvid|Smith et al.|2006}} }}
{{refend}}

== External links ==
*[https://web.archive.org/web/20001101213011/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?1221 Germplasm Resources Information Network: Blechnaceae]

{{Plant classification}}
{{Fern classification}}
{{Taxonbar|from=Q932153}}

[[Category:Blechnaceae| ]]
[[Category:Fern families]]
[[Category:Taxa named by Edward Newman]]