Dryopteridaceae

The Dryopteridaceae are a family of leptosporangiate ferns in the order Polypodiales. They are known colloquially as the wood ferns. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family is placed in the suborder Polypodiineae. Alternatively, it may be treated as the subfamily Dryopteridoideae of a very broadly defined family Polypodiaceae '' sensu lato''.

The family contains about 1700 species and has a cosmopolitan distribution. Species may be terrestrial, epipetric, hemiepiphytic, or epiphytic. Many are cultivated as ornamental plants. The largest genera are ''Elaphoglossum'' (600+), ''Polystichum'' (260), ''Dryopteris'' (225), and ''Ctenitis'' (150). These four genera contain about 70% of the species. Dryopteridaceae diverged from the other families in eupolypods I about 100 million years ago.

Description

The rhizomes are often stout, creeping, ascending, or erect, and sometimes scandent or climbing, with non clathrate scales at apices. Fronds 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 areoles 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 or peltate. Sporangia have three-rowed, short to long stalks; spores arereniform, Spore#Anatomy, monolete, spore perine, perine or Glossary of botanical terms#W, winged.

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. 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. 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. Some authors have already treated these genera as outside of the Dryopteridaceae.

In 2007, a phylogenetic study of DNA sequences showed that ''Pleocnemia'' should be transferred from the Tectariaceae to the Dryopteridaceae. In 2010, in a paper on bolbitidoid ferns, ''Arthrobotrya'' was resurrected from ''Teratophyllum''. Later that year, ''Mickelia'' was described as a new genus.

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.

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

The Dryopteridaceae Wilhelm Gustav Franz Herter, Herter, under the classification system of Christenhusz and Chase (2014), were submerged as subfamily Dryopteridoideae Johann Heinrich Friedrich Link, Link, one of eight subfamilies constituting family Polypodiaceae. This family corresponds to the clade eupolypods I. The Pteridophyte Phylogeny Group classification of 2016 (PPG I) retained the family.

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), shows a likely phylogenetics, phylogenetic relationship between Dryopteridaceae and the other families of the clade.

Subdivision

The PPG I classification divides the family into three subfamilies, listed below.
* Subfamily Polybotryoideae H.M.Liu & X.C.Zhang
** ''Cyclodium'' C.Presl
** ''Maxonia'' C.Chr.
** ''Olfersia (plant), Olfersia'' Raddi
** ''Polybotrya'' Humb. & Bonpl. ex Willd.
** ''Polystichopsis'' (J.Sm.) Holttum
** ''Stigmatopteris'' C.Chr.
** ''Trichoneuron'' Ching
* Subfamily Elaphoglossoideae (Pic.Serm.) Crabbe, Jermy & Mickel
** ''Arthrobotrya'' J.Sm.
** ''Bolbitis'' Schott
** ''Elaphoglossum'' Schott ex J.Sm.
** ''Lastreopsis'' Ching
** ''Lomagramma'' J.Sm.
** ''Megalastrum'' Holttum
** ''Mickelia'' R.C.Moran, Labiak & Sundue
** ''Parapolystichum'' (Keyserl.) Ching
** ''Pleocnemia'' C.Presl
** ''Rumohra'' Raddi
** ''Teratophyllum'' Mett. ex Kuhn
* Subfamily Dryopteridoideae Link
** ''Arachniodes'' Blume
** ''Ctenitis'' (C.Chr.) C.Chr.
** ''Cyrtomium'' C.Presl
** ''Dryopteris'' Adans.
** ''Phanerophlebia (plant), Phanerophlebia'' C.Presl
** ''Polystichum'' Roth

''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). ''Dryopolystichum'' has been placed in Lomariopsidaceae.

References

Bibliography

*

External links

''Phytotaxa''

{{Authority control

Dryopteridaceae,
Fern families