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Blechnaceae is a family of
fern The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...
s in the order
Polypodiales The Order (biology), order Polypodiales encompasses the major lineages of polypod ferns, which comprise more than 80% of today's fern species. They are found in many parts of the world including Tropics, tropical, semitropical and Temperate clima ...
, with a
cosmopolitan distribution In biogeography, a cosmopolitan distribution is the range of a taxon that extends across most or all of the surface of the Earth, in appropriate habitats; most cosmopolitan species are known to be highly adaptable to a range of climatic and en ...
. Its status as a family and the number of genera included have both varied considerably. In the
Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group (PPG) is an informal international group of systematic botanists who collaborate to establish on the classification of pteridophytes ( lycophytes and ferns) that reflects knowledge about plant relationships discove ...
classification of 2016 (PPG I), the family has 24 genera, and excludes genera placed in the separate family
Onocleaceae Onocleaceae is a small family of terrestrial ferns in the order Polypodiales. It is placed in the suborder Aspleniineae in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Alternatively, the family, along with Blechnaceae, may be ...
. The family is divided into three subfamilies, including Blechnoideae s.s. Alternatively, the entire family may be treated as the subfamily Blechnoideae s.l. of a very broadly defined family
Aspleniaceae The Aspleniaceae (spleenworts) are a family of ferns, included in the order Polypodiales. The composition and classification of the family have been subject to considerable changes. In particular, there is a narrow circumscription, Aspleniaceae ...
, and include genera others place in Onocleaceae.


Description

Most are ground dwelling, some are climbers, such as ''
Stenochlaena ''Stenochlaena'' is a genus of ferns of the plant family Blechnaceae. Six species were formally accepted in an April 2013 scientific review of the genus, first written some years earlier and submitted in 2009. One additional species ''S. hai ...
''. A characteristic feature of many species is that the young opening
frond A frond is a large, divided leaf. In both common usage and botanical nomenclature, the leaves of ferns are referred to as fronds and some botanists restrict the term to this group. Other botanists allow the term frond to also apply to the lar ...
s are usually tinged with red.


Taxonomy

The family was created by
Newman Newman is a surname of Germanic Anglo-Saxon origins. Newman is the modern English form of the name used in Great Britain and among people of British ancestry around the world (as is 'Numan'), while Neumann (with variant spellings) is used in Ger ...
in 1844. In 2014, Christenhusz and Chase submerged it as subfamily Blechnoideae within the family
Aspleniaceae The Aspleniaceae (spleenworts) are a family of ferns, included in the order Polypodiales. The composition and classification of the family have been subject to considerable changes. In particular, there is a narrow circumscription, Aspleniaceae ...
and included
Onocleaceae Onocleaceae is a small family of terrestrial ferns in the order Polypodiales. It is placed in the suborder Aspleniineae in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Alternatively, the family, along with Blechnaceae, may be ...
in it. The
PPG I The Pteridophyte Phylogeny Group (PPG) is an informal international group of systematic botanists who collaborate to establish on the classification of pteridophytes ( lycophytes and ferns) that reflects knowledge about plant relationships discove ...
classification of 2016 restored it to family status. Blechnaceae is a member of the eupolypods II clade (now the suborder Aspleniineae), in the order
Polypodiales The Order (biology), order Polypodiales encompasses the major lineages of polypod ferns, which comprise more than 80% of today's fern species. They are found in many parts of the world including Tropics, tropical, semitropical and Temperate clima ...
. It is related to other families in the clade as in the following 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 ''Woodwardia'' is a genus of ferns in the family Blechnaceae, in the suborder Aspleniineae (eupolypods II) of the order Polypodiales. Species are known as netted-chain ferns. The genus is native to warm temperate and subtropical regions of the ...
'', sister to all other species that can be treated as the single genus ''
Blechnum ''Blechnum'', known as hard fern, is a genus of ferns in the family Blechnaceae, subfamily Blechnoideae, according to the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Two very different circumscriptions of the genus are used by ...
''. However, the subclade sister to the rest of ''Blechnum'' ''sensu lato'' contains the vining taxa ''Stenochlaena'', ''
Salpichlaena ''Salpichlaena'' is a genus containing two species of fern in the family Blechnaceae. Species of the genus are native to Central and South America. These ferns have climbing leaves with a rachis that twines around tree branches and other support ...
'' J.Sm. 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 ''Brainea'' is a genus of ferns in the family Blechnaceae, subfamily Blechnoideae, with a single species ''Brainea insignis'', according to the Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group (PPG) is an informal international gro ...
'', ''
Doodia ''Doodia'' is a genus of ferns in the family Blechnaceae, subfamily Blechnoideae, in the suborder Aspleniineae (eupolypods II). It is named after Samuel Doody (1656–1706), an English botanist. Distribution of the genus includes parts of Austra ...
'', '' Pteridoblechnum'' and ''
Sadleria ''Sadleria'' is a genus of six species of ferns in the family Blechnaceae, all endemic to Hawaii.Fern pages''Sadleria''/ref> Taxonomy Georg Friedrich Kaulfuss distinguished the genus in 1824 based on samples of ''S. cyatheoides'' acquired by ...
'' belong to ''Blechnum'' sensu lato." Perrie et al. (2014) retained the family rank and excluded '' Onoclea'' as a separate family,
Onocleaceae Onocleaceae is a small family of terrestrial ferns in the order Polypodiales. It is placed in the suborder Aspleniineae in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Alternatively, the family, along with Blechnaceae, may be ...
, 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 In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
. They did not consider ''
Blechnum ''Blechnum'', known as hard fern, is a genus of ferns in the family Blechnaceae, subfamily Blechnoideae, according to the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Two very different circumscriptions of the genus are used by ...
'' as monophyletic and recommended revision of intergeneric boundaries, resulting in seven genera. 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 A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...
nature of ''Blechnum'' was to create a series of monophyletic segregate genera, resulting in 24 genera in total. The relationship between the three subfamilies was found to be as follows: The approximate relationship between some of the taxa used in the classifications of Christenhusz and Chase (2014) and PPG I (2016) is shown in the table below.


Subfamilies and genera

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. Subfamily Woodwardioideae Gasper, V.A.O.Dittrich & Salino *''
Anchistea ''Anchistea'' is a genus of Leptosporangiate fern, leptosporangiate ferns in the family Blechnaceae. It has only one species, ''Anchistea virginica'' (Synonym (taxonomy), synonym ''Woodwardia virginica'') the Virginia chain fern, which has long c ...
'' C.Presl *''
Lorinseria ''Lorinseria'' is a genus of fern in the subfamily Woodwardioideae of the family Blechnaceae. Its only species is ''Lorinseria areolata'' (synonym ''Woodwardia areolata''), the netted chain fern, native to eastern North America. The monotypic g ...
'' C.Presl *''
Woodwardia ''Woodwardia'' is a genus of ferns in the family Blechnaceae, in the suborder Aspleniineae (eupolypods II) of the order Polypodiales. Species are known as netted-chain ferns. The genus is native to warm temperate and subtropical regions of the ...
'' Sm. Subfamily Stenochlaenoideae (Ching) J.P.Roux *''
Salpichlaena ''Salpichlaena'' is a genus containing two species of fern in the family Blechnaceae. Species of the genus are native to Central and South America. These ferns have climbing leaves with a rachis that twines around tree branches and other support ...
'' J.Sm. *''
Stenochlaena ''Stenochlaena'' is a genus of ferns of the plant family Blechnaceae. Six species were formally accepted in an April 2013 scientific review of the genus, first written some years earlier and submitted in 2009. One additional species ''S. hai ...
'' J.Sm. *''
Telmatoblechnum ''Telmatoblechnum'' is a genus of ferns in the family Blechnaceae, subfamily Stenochlaenoideae. Species , the ''Checklist of Ferns and Lycophytes of the World'' and ''Plants of the World Online'' accepted two species: *'' Telmatoblechnum indicum ...
'' Perrie, D.J.Ohlsen & Brownsey Subfamily Blechnoideae Gasper, V.A.O.Dittrich & Salino *'' Austroblechnum'' Gasper & V.A.O.Dittrich *''
Blechnidium ''Blechnidium'' is a genus of ferns in the family Blechnaceae, subfamily Blechnoideae, with a single species ''Blechnidium melanopus'', according to the Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group (PPG) is an informal interna ...
'' T.Moore *'' Blechnopsis'' C.Presl *''
Blechnum ''Blechnum'', known as hard fern, is a genus of ferns in the family Blechnaceae, subfamily Blechnoideae, according to the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Two very different circumscriptions of the genus are used by ...
'' L. *''
Brainea ''Brainea'' is a genus of ferns in the family Blechnaceae, subfamily Blechnoideae, with a single species ''Brainea insignis'', according to the Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group (PPG) is an informal international gro ...
'' J.Sm. *'' Cleistoblechnum'' Gasper & Salino *''
Cranfillia ''Cranfillia'' is a genus of ferns in the family Blechnaceae, subfamily Blechnoideae, according to the Pteridophyte Phylogeny Group classification of 2016 (PPG I). The genus is accepted in a 2016 classification of the family Blechnaceae, but othe ...
'' Gasper & V.A.O.Dittrich *'' Diploblechnum'' Hayata *''
Doodia ''Doodia'' is a genus of ferns in the family Blechnaceae, subfamily Blechnoideae, in the suborder Aspleniineae (eupolypods II). It is named after Samuel Doody (1656–1706), an English botanist. Distribution of the genus includes parts of Austra ...
'' R.Br. *''
Icarus In Greek mythology, Icarus (; , ) was the son of the master craftsman Daedalus, the architect of the labyrinth of Crete. After Theseus, king of Athens and enemy of King Minos, escaped from the labyrinth, Minos suspected that Icarus and Daedalu ...
'' Gasper & Salino *'' Lomaria'' Willd. *'' Lomaridium'' C.Presl *'' Lomariocycas'' (J.Sm.) Gasper & A.R.Sm. *'' Neoblechnum'' Gasper & V.A.O.Dittrich *×'' Oceanidoodia'' Liu, Schuettp. & H. Schneid. *'' Oceaniopteris'' Gasper & Salino *'' Parablechnum'' C.Presl *''
Sadleria ''Sadleria'' is a genus of six species of ferns in the family Blechnaceae, all endemic to Hawaii.Fern pages''Sadleria''/ref> Taxonomy Georg Friedrich Kaulfuss distinguished the genus in 1824 based on samples of ''S. cyatheoides'' acquired by ...
'' Kaulf. *''
Struthiopteris ''Struthiopteris'' is a genus of ferns belonging to the family Blechnaceae, subfamily Blechnoideae. Species In the circumscription used in the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the genus contains five species. The ''Ch ...
'' Scop. Subfamily ''incertae sedis'' *'' Trawetsia''
Ypresian In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
,
Princeton Chert The Princeton Chert is a fossil locality in British Columbia, Canada, which comprises an anatomically preserved flora of Eocene Epoch age, with rich species abundance and diversity. It is located in exposures of the Allenby Formation on the east ...


References


Bibliography

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External links


Germplasm Resources Information Network: Blechnaceae
{{Taxonbar, from=Q932153 Fern families Taxa named by Edward Newman