Hymenophyllaceae

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Hymenophyllaceae
Temporal range: Upper Triassic–Recent
Hymenophyllum tunbrigense in Luxembourg
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Division: Polypodiophyta
Class: Polypodiopsida
Subclass: Polypodiidae
Order: Hymenophyllales
A.B.Frank
Family: Hymenophyllaceae
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Genera

See text

The Hymenophyllaceae, the filmy ferns and bristle ferns, are a family of two to nine genera (depending on classification system) and about 650 known species

Upper Triassic.[2]

Description

They often appear as very dark green or even black clumps and may be mistaken for a robust

trilete spores. The spores grow into thread- or ribbon-like gametophytes; in many species, the gametophyte has an extended, independent lifespan and can reproduce asexually by fragmenting or releasing gemmae.[3]

Individual plants may persist for many years.

Taxonomy

In the molecular phylogenetic classification of Smith et al. in 2006, the Hymenophyllales, containing the single family Hymenophyllaceae, were placed in class Polypodiopsida sensu stricto (the leptosporangiate ferns).[3] The linear sequence of Christenhusz et al. (2011), intended for compatibility with the classification of Chase and Reveal (2009)[4] which placed all land plants in Equisetopsida,[5] reclassified Smith's Polypodiopsida as subclass Polypodiidae and placed the Hymenophyllales there. The circumscription of the order and its families was not changed,[4] and that circumscription and placement in Polypodiidae has subsequently been followed in the classifications of Christenhusz and Chase (2014)[6] and PPG I (2016).[7]

The division of the family into genera was disputed, as of October 2019. Traditionally, only two genera of Hymenophyllaceae have been recognized: (1)

monophyletic clades of fairly equal size, but they are only roughly aligned with the two traditional genera. For example, the traditional Trichomanes subtaxa Pleuromanes and Cardiomanes were shown to belong to the "hymenophylloid" clade. To reflect these recent discoveries Atsushi Ebihara and Kunio Iwatsuki, in 2006, revised the taxonomy of Hymenophyllaceae to place all species of the "hymenophylloid" clade in a single genus Hymenophyllum, and to place the eight clear "trichomanoid" subclades in eight corresponding genera.[8]

This subdivision was recognized by Smith et al. in 2006[3] and Christenhusz et al. in 2011,[4] but Christenhusz and Chase, in 2014, reverted to combining the trichomanoid clades into Trichomanes.[6] The PPG I classification of 2016 again recognizes the segregate genera (and treats the two clades as subfamilies, Hymenophylloideae and Trichomanoideae),[7] although the segregate genera are not always accepted by contemporary floras; e.g., as of 2016, the Flora of New Zealand preferred to recognize Trichomanes s.l. due to the difficulty of morphologically distinguishing the segregate genera.[9]

Genera

The genera used in PPG I and the subgenera assigned by the system of Ebihara et al. are:

Phylogeny of Hymenophyllaceae[8]
Phylogeny of Hymenophyllaceae[10][11]
Hymenophylloideae

Hymenophyllum

Trichomanoideae
  • Hymenophylloideae (the "hymenophylloid" clade):
    • Hymenophyllum Sm. 1793 – about 250 species
      • subg. Hymenophyllum – about 100 species
      • subg. Sphaerocionium (C.Presl) C.Chr. 1934 – about 70 species
      • subg. Mecodium C.Presl ex Copel. 1937 – more than 35 species
      • subg. Globosa (Prantl) Ebihara & K.Iwats. 2006 – about 25 species
      • subg. Pleuromanes (C.Presl) Ebihara & K.Iwats. 2006 – 5 species
      • subg. Myrmecostylum (C.Presl) Ebihara & K. Iwats. 2006 – at least 8 species
      • subg. Hymenoglossum (C.Presl) R.M.Tryon & A.F.Tryon 1981 – at least 3 species
      • subg. Fuciformia Ebihara & K.Iwats. 2006 – 2 species
      • subg. Diploöphyllum (Bosch) Ebihara & K.Iwats. 2006 – 1 species
      • subg. Cardiomanes (C. Presl) Ebihara & K.Iwats. 2006 – 1 species
  • Trichomanoideae (the "trichomanoid" clade) (sometimes all included in a single broad genus Trichomanes with about 400 species):
    • Didymoglossum Desv. 1827 – more than 30 species
      • subg. Didymoglossum – more than 20 species
      • subg. Microgonium (C.Presl) Ebihara & K.Iwats. 2006 – more than 10 species
    • Crepidomanes (C.Presl) C.Presl 1849 – more than 30 species
      • subg. Crepidomanes
      • subg. Nesopteris (Copel.) Ebihara & K.Iwats. 2006
    • Polyphlebium Copel. 1938 – about 15 species
    • Vandenboschia Copel. 1938 – more than 15 species
      • subg. Vandenboschia – more than 15 species
      • subg. Lacosteopsis (Prantl) Ebihara & K.Iwats. 2006 – at least 2 species
    • Abrodictyum C.Presl 1843 – about 25 species
      • subg. Abrodictyum – about 15 species
      • subg. Pachychaetum (C.Presl) Ebihara & K.Iwats. 2006 – more than 10 species
    • Trichomanes L. 1753 – more than 60 species
      • subg. Afrotrichomanes Dubuisson & Ebihara 2021
      • subg. Davalliopsis (Bosch) Ebihara & K.Iwats. 2006 – at least 1 species
      • subg. Feea (Bory) Hook. 1844 – more than 5 species
      • subg. Lacostea (Bosch) C. Chr. 1906 – more than 4 species
      • subg. Trichomanes – more than 30 species
    • Cephalomanes C.Presl 1843 – about 4 species
    • Callistopteris Copel. 1938 – about 5 species

Distribution and habitat

The great majority of the species are found in

Faeroes and South Iceland, while in North America, they are restricted (often occurring solely as gametophytes
) to the humid eastern third of the continent and the rainforests of the Pacific Northwest.

References