The Athyriaceae (ladyferns and allies) are a family of terrestrial

fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except ...

s in the order

Polypodiales The 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 tropical, semitropical and temperate areas. Description Polypodiale ...

. In the

Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group, or PPG, is an informal international group of systematic botanists who collaborate to establish a consensus on the classification of pteridophytes (lycophytes and ferns) that reflects knowledge about plant relatio ...

classification of 2016 (PPG I), the family is placed in the suborder

Aspleniineae Aspleniineae is a suborder of ferns in the order Polypodiales. It is equivalent to the clade eupolypods II in earlier systems; it is also treated as a single very broadly defined family Aspleniaceae. The suborder generally corresponds with the ...

, and includes two genera. Alternatively, it may be treated as the subfamily Athyrioideae 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 ...

. The family has with a cosmopolitan distribution.

Description

Species of the Athyriaceae are terrestrial or

lithophytic Lithophytes are plants that grow in or on rocks. They can be classified as either epilithic (or epipetric) or endolithic; epilithic lithophytes grow on the surfaces of rocks, while endolithic lithophytes grow in the crevices of rocks (and are als ...

, less commonly aquatic. They grow from various kinds of rhizome: short or long, creeping or erect, branched or not. The distribution and evolution of

characters Character or Characters may refer to: Arts, entertainment, and media Literature * ''Character'' (novel), a 1936 Dutch novel by Ferdinand Bordewijk * ''Characters'' (Theophrastus), a classical Greek set of character sketches attributed to The ...

in the family is complex, and the genera have few constant features by which they can be identified. The sporangia have stalks two or three cells wide in the middle, and contain brown

monolete In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae ...

spores.

Taxonomy

Earlier classifications

The family was first created by Arthur H.G. Alston in 1956. It has had a varied history. In 2014, Christenhusz and Chase submerged it as the subfamily Athyrioideae within the family

, a status maintained by

Plants of the World Online Plants of the World Online (POWO) is an online database published by the Royal Botanic Gardens, Kew. It was launched in March 2017 with the ultimate aim being "to enable users to access information on all the world's known seed-bearing plants ...

. The PPG I classification of 2016 restored it to family status. Athyriaceae is a member of the eupolypods II clade (now the suborder Aspleniineae), in the order

. It is related to other families in the clade as in the following cladogram: The Athyriaceae in the past included ''

Cystopteris ''Cystopteris'' is a genus of ferns in the family Cystopteridaceae. These are known generally as bladderferns or fragile ferns. They grow in temperate areas worldwide. This is a very diverse genus and within a species individuals can look quite ...

'' and ''

Gymnocarpium ''Gymnocarpium'' is a small genus of ferns, called oak ferns. It was once placed with various other groups, including the dryopteroid ferns and the athyrioid ferns. Cladistic analysis has demonstrated that ''Gymnocarpium'' and ''Cystopteris'' ...

'' (now part of

Dennstaedtiaceae Dennstaedtiaceae is one of fifteen families in the order Polypodiales, the most derived families within monilophytes (ferns). It comprises 10 genera with ca 240 known species, including one of the world's most abundant ferns, ''Pteridium aquilin ...

). The family has been subsumed in the family

Woodsiaceae ''Woodsia'' is a genus of ferns in the order Polypodiales. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), it is the only genus in the family Woodsiaceae, placed in the suborder Aspleniineae. The family can also be trea ...

, but a Woodsiaceae defined in this way may be

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

if it omits the Onocleaceae and Blechnaceae (as of 2006, the evidence was not clear).

Genera

circumscribed In geometry, the circumscribed circle or circumcircle of a polygon is a circle that passes through all the vertices of the polygon. The center of this circle is called the circumcenter and its radius is called the circumradius. Not every po ...

in PPG I, Athyriacae contains the following genera. *''

Athyrium ''Athyrium'' (lady-fern) is a genus of about 180 species of terrestrial ferns, with a cosmopolitan distribution. It is placed in the family Athyriaceae, in the order Polypodiales. Its genus name is from Greek '' a-'' ('without') and Latinized ...

'' Roth (including ''Anisocampium'' and ''Cornopteris'' Nakai) *'' Deparia'' Hook. & Grev. *''

Diplazium ''Diplazium'' is a genus of ferns that specifically includes the approximately 400 known species of twinsorus ferns. The Greek root is ''diplazein'' meaning ''double'': the indusia in this genus lie on both sides of the vein. These ferns were e ...

'' Sw. , the ''Checklist of Ferns and Lycophytes of the World'' recognizes three further genera, which other sources include in ''Athyrium'': *''Anisocampium'' C.Presl *''Cornopteris'' Nakai *''Pseudathyrium'' Newman The genera have the following

phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

relationship:

Distribution and habitat

Athyriaceae has a worldwide distribution, particularly the genus ''Athyrium''. Most species of Athyriaceae are medium-sized terrestrial ferns, growing in the

understorey In forestry and ecology, understory (American English), or understorey (Commonwealth English), also known as underbrush or undergrowth, includes plant life growing beneath the forest canopy without penetrating it to any great extent, but abov ...

below trees and shrubs.

References

{{Taxonbar, from=Q3774137 Fern families