Thelypteridaceae is a family of about 900 species 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 ...

. 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), it 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 o ...

. Alternatively, the family may be submerged in a very broadly defined family Aspleniaceae as the subfamily Thelypteridoideae. The ferns are

terrestrial Terrestrial refers to things related to land or the planet Earth, as opposed to extraterrestrial. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on o ...

, with the exception of a few which are

lithophyte 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 ...

s (grow on rocks). The bulk of the species are tropical, although there are a number of temperate species. These ferns typically have creeping

rhizome In botany and dendrology, a rhizome ( ) is a modified subterranean plant stem that sends out roots and Shoot (botany), shoots from its Node (botany), nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from ...

s. The

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 simply

pinnate Pinnation (also called pennation) is the arrangement of feather-like or multi-divided features arising from both sides of a common axis. Pinnation occurs in biological morphology, in crystals, such as some forms of ice or metal crystals, and ...

to pinnate-

pinnatifid Pinnation (also called pennation) is the arrangement of feather-like or multi-divided features arising from both sides of a common axis. Pinnation occurs in biological morphology, in crystals, such as some forms of ice or metal crystals, and in ...

. There is either no frond dimorphism or only mild dimorphism, either open

venation Venation may refer to: * Venation (botany), the arrangement of veins in leaves * Wing venation, the arrangement of veins in insect wings See also * * Vernation Vernation or leafing is the formation of new leaves or fronds. In plant anatomy, it ...

or very simple

anastomosing An anastomosis (, : anastomoses) is a connection or opening between two things (especially cavities or passages) that are normally diverging or branching, such as between blood vessels, leaf veins, or streams. Such a connection may be normal (suc ...

. The sori are mostly reniform in shape and have indusia, except for the ''Phegopteris'' group.

Classification

During the early and mid 1900's all thelypterioid ferns were included in the genus ''

Dryopteris :''The moth genus ''Dryopteris'' is now considered a junior synonym of ''Oreta. ''Dryopteris'' , commonly called the wood ferns, male ferns (referring in particular to ''Dryopteris filix-mas''), or buckler ferns, is a fern genus in the family Dry ...

'' because of the sorus shape. However, there are a great many differences between the groups, and these plants are now segregated in their own family. Genetic evidence shows that the family is clearly

monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...

Phylogenetic relationships

The following cladogram for the suborder

(as eupolypods II), based on Lehtonen (2011), and Rothfels & al. (2012), shows a likely

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

relationship between the Thelypteridaceae and the other families of the clade.

Genera

The family can be divided into two major clades, which the

classification of 2016 (PPG I) recognizes as two subfamilies, Phegopteridoideae and Thelypteroideae. Their division into genera has been described as "highly controversial and fluctuating". The family includes several complexes of species that are difficult to distinguish, and seem to represent a remarkable evolutionary radiation. Some researchers include the entire family Thelypteridaceae in the genus ''

Thelypteris ''Thelypteris'' (maiden ferns) is a genus of ferns in the subfamily Thelypteridoideae, family Thelypteridaceae, order Polypodiales. Two radically different circumscriptions of the genus are in use . In the Pteridophyte Phylogeny Group classifi ...

''; others divide the family into as many as 30 genera. An intermediate position is to place the bulk in ''Thelypteris'' (which can then be divided into subgenera and sections corresponding to the genera of other authors) but to separate out ''

Phegopteris ''Phegopteris'' is a genus of ferns in the family Thelypteridaceae, subfamily Phegopteridoideae, in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). They are known collectively as the beech ferns. Species are native to Asia, North ...

'' and ''

Macrothelypteris ''Macrothelypteris'' is a genus of ferns in the family Thelypteridaceae, subfamily Phegopteridoideae, in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). The genus name means 'large ''Thelypteris. Species , the ''Checklist of ...

''. Another choice is to divide the family into a half a dozen or so genera. The Pteridophyte Phylogeny Group classification of 2016 (PPG I) accepts 30 genera: *Subfamily Phegopteridoideae Salino, A.R.Sm. & T.E.Almeid **''

'' (H.Ito) Ching **''

'' (C.Presl) Fée **''

Pseudophegopteris ''Phegopteris'' is a genus of ferns in the family Thelypteridaceae, subfamily Phegopteridoideae, in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). They are known collectively as the beech ferns. Species are native to Asia, North ...

'' Ching *Subfamily Thelypteridoideae C.F.Reed **'' Steiropteris'' (C.Chr.) Pic.Serm. **Tribe Thelypterideae Ching 1963 ***''

'' Schmidel **Tribe Amauropelteae Wei, Liu & Liu ***''

Amauropelta ''Amauropelta'' is a Monophyly, monophyletic genus of fern, the largest in the Thelypteridaceae family, with 233 species.Fawcett, S. & A.R. Smith. 2021. A generic classification of the Thelypteridaceae. Sida, Bot. Misc. 59. BRIT Press, Fort Wor ...

'' Kunze ***''

Coryphopteris ''Coryphopteris'' is a genus of ferns in the family Thelypteridaceae, subfamily Thelypteridoideae, in the Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group (PPG) is an informal international group of systematic botanists who collabo ...

'' Holttum ***'' Metathelypteris'' (H.Ito) Ching **Tribe Oreopterideae Wei, Liu & Liu ***''

Oreopteris ''Oreopteris'' is a genus of ferns belonging to the family Thelypteridaceae. The species of this genus are found in Eurasia and North America. Species: * ''Oreopteris elwesii'' (Baker ex Hook. & Baker) Holttum * ''Oreopteris limbosperma ''O ...

'' Holub **Tribe Leptogrammeae Todaro 1866 ***'' Cyclogramma'' Tagawa ***'' Leptogramma'' Smith ***'' Stegnogramma'' Blume **Tribe Meniscieae Fée 1850-1852 ex Pfeiffer 1873 ***Subtribe Goniopteridinae Ching 1963 ****'' Goniopteris'' C.Presl ***Subtribe Menisciinae Payer 1850 ****'' Ampelopteris'' Kunze ****''

Cyclosorus ''Cyclosorus'' is a genus of ferns in the family Thelypteridaceae, subfamily Thelypteridoideae, in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Other sources sink ''Cyclosorus'' into a very broadly defined genus ''Thelypteri ...

'' Link ****'' Meniscium'' Schreb. ****'' Mesophlebion'' Holttum ***Subtribe Pseudocyclosorinae Ching 1963 ****'' Abacopteris'' Fée ****'' Amblovenatum'' J.P.Roux ****'' Chingia'' Holttum ****'' Christella'' H.Lév. ****'' Glaphyropteridopsis'' Ching ****'' Grypothrix'' (Holttum 1971) Fawcett & Smith ****'' Menisciopsis'' (Holttum 1982) Fawcett & Smith ****'' Menisorus'' Alston ****'' Mesopteris'' Ching ****'' Pakau'' Fawcett & Smith ****'' Parathelypteris'' (H.Ito) Ching ****'' Pelazoneuron'' (Holttum 1974) Smith & Fawcett ****'' Plesioneuron'' (Holttum) Holttum ****''

Pneumatopteris ''Pneumatopteris'' is a genus of about 80 species of Terrestrial plant, terrestrial ferns in the Family (biology), family Thelypteridaceae. The range of the genus extends from tropical Africa, through Asia, Malesia and Australia to the List of i ...

'' Nakai ****'' Pronephrium'' C.Presl 'Nannothelypteris'' Holttum">Nannothelypteris.html" ;"title="'Nannothelypteris">'Nannothelypteris'' Holttum****''Pseudocyclosorus'' Ching ****''Reholttumia'' Fawcett & Smith ****''Sphaerostephanos'' J.Sm. ****''Strophocaulon'' Fawcett & Smith ****''Trigonospora'' Holttum Some species of ''Pronephrium'' were split off into other genera in 2021 as a result of a Phylogenetics, phylogenetic study of the family Thelypteridaceae: *'' Abacopteris'' Fée *'' Grypothrix'' (Holttum) S.E.Fawc. & A.R.Sm. *'' Menisciopsis'' (Holttum) S.E.Fawc. & A.R.Sm The extinct genus '' Holttumopteris'' from the

Cenomanian The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...

aged

Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. Th ...

has been suggested to have affinities with the family, but several important diagnostic characters are not preserved.

References

{{Authority control Fern families