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[1] of ferns, with a subcosmopolitan distribution, but generally restricted to very damp places or to locations where they are wetted by spray from waterfalls or springs. Fossil evidence shows that ferns of the family Hymenophyllaceae have existed since at least the Upper Triassic.[2]
| Hymenophyllaceae Temporal range:
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| Hymenophyllum tunbrigense in Luxembourg | |
Scientific classification 
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| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Division: | Polypodiophyta |
| Class: | Polypodiopsida |
| Subclass: | Polypodiidae |
| Order: | Hymenophyllales A.B.Frank |
| Family: | Hymenophyllaceae Link |
| Genera | |
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See text | |
Description
editThey often appear as very dark green or even black clumps and may be mistaken for a robust moss or liverwort. The rhizome is usually thin and wiry and the fronds variously pinnate with a single strand ("nerve") of vascular tissue. As in most ferns, young fronds have circinate vernation. In most species, the frond, apart from the vascular tissue, is only a single cell thick, and they do not have any stomata. The cuticle is also greatly reduced or absent,[3] leaving filmy ferns poikilohydric and very susceptible to desiccation where a reliable water supply is not present.[4] The leaves occasionally bear hairs, but scales are generally not present.[3] The sori are borne at the leaf margins[3] at the end of the nerve. They are protected by conical, bivalvate, or tubular indusia. Within the sori, sporangia mature starting at the apex of the sorus and progressing to the base. They have a continuous, oblique annulus and release round, green 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
editIn 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)[5] which placed all land plants in Equisetopsida,[6] reclassified Smith's Polypodiopsida as subclass Polypodiidae and placed the Hymenophyllales there. The circumscription of the order and its families was not changed,[5] and that circumscription and placement in Polypodiidae has subsequently been followed in the classifications of Christenhusz and Chase (2014)[7] and PPG I (2016).[8]
The division of the family into genera was disputed, as of October 2019[update]. Traditionally, only two genera of Hymenophyllaceae have been recognized: (1) Hymenophyllum with bivalved involucres, and (2) Trichomanes s.l. with tubular involucres. Subsequent proposals have created 34 genera (Copeland 1938), 6 genera (Morton 1968), 47 genera (Sermolli 1977), and 8 genera (Iwatsuki 1984). These classifications all had only limited regional acceptance. Recent molecular phylogenic studies do show two distinct 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.[9]
This subdivision was recognized by Smith et al. in 2006[3] and Christenhusz et al. in 2011,[5] but Christenhusz and Chase, in 2014, reverted to combining the trichomanoid clades into Trichomanes.[7] The PPG I classification of 2016 again recognizes the segregate genera (and treats the two clades as subfamilies, Hymenophylloideae and Trichomanoideae),[8] 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.[10]
Genera
editThe genera used in PPG I and the subgenera assigned by the system of Ebihara et al. are:
| Phylogeny of Hymenophyllaceae[9] | |||||||||
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| Phylogeny of Hymenophyllaceae[11][12] | |||||||||||||||
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- 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
- Hymenophyllum Sm. 1793 – about 250 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
- Didymoglossum Desv. 1827 – more than 30 species
Distribution and habitat
editThe great majority of the species are found in tropical rainforests, but some also occur in temperate rainforests (particularly New Zealand, with 25 species) and slightly drier forest regions. In Europe they are restricted to the Atlantic Ocean fringes of the continent, notably in the Azores, Ireland, and western Great Britain, but one species (Hymenophyllum tunbrigense) locally east to Luxembourg, another (H. wilsonii) so far north as West Norway, 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
edit- ^ Christenhusz, M. J. M.; Byng, J. W. (2016). "The number of known plants species in the world and its annual increase". Phytotaxa. 261 (3). Magnolia Press: 201–217. doi:10.11646/phytotaxa.261.3.1.
- ^ A filmy fern from the Upper Triassic of North Carolina (USA) - Axsmith et al. 88 (9): 1558 - American Journal of Botany
- ^ a b c d e f Smith, Alan R.; Pryer, Kathleen M.; Schuettpelz, Eric; Korall, Petra; Schneider, Harald; Wolf, Paul G. (August 2006). "A classification for extant ferns" (PDF). Taxon. 55 (3): 705–731. doi:10.2307/25065646. JSTOR 25065646. Archived from the original (PDF) on 2021-12-09. Retrieved 2016-12-22.
- ^ Light and desiccation responses of some Hymenophyllaceae (filmy ferns) from Trinidad, Venezuela and New Zealand: poikilohydry in a light-limited but low evaporation ecological niche
- ^ a b c Christenhusz, Maarten J. M.; Zhang, Xian-Chun; Schneider, Harald (18 February 2011). "A linear sequence of extant families and genera of lycophytes and ferns" (PDF). Phytotaxa. 19: 7–54. doi:10.11646/phytotaxa.19.1.2.
- ^ Chase, Mark W.; Reveal, James L. (October 2009). "A phylogenetic classification of the land plants to accompany APG III". Botanical Journal of the Linnean Society. 161 (2): 122–127. doi:10.1111/j.1095-8339.2009.01002.x.
- ^ a b Christenhusz, Maarten J. M.; Chase, Mark W. (13 February 2014). "Trends and concepts in fern classification". Annals of Botany. 113 (4): 571–594. doi:10.1093/aob/mct299. PMC 3936591. PMID 24532607.
- ^ a b The Pteridophyte Phylogeny Group (November 2016). "A community-derived classification for extant lycophytes and ferns". Journal of Systematics and Evolution. 54 (6): 563–603. doi:10.1111/jse.12229. S2CID 39980610.
- ^ a b Ebihara, Atsushi; Dubuisson, Jean-Yves; Iwatsuki, Kunio; Hennequin, Sabine; Ito, Motomi (July 2006). "A Taxonomic Revision of Hymenophyllaceae". Blumea. 51 (2): 221–280. doi:10.3767/000651906x622210.
- ^ "Hymenophyllaceae Mart". Flora of New Zealand. Retrieved 22 December 2016.
- ^ Nitta, Joel H.; Schuettpelz, Eric; Ramírez-Barahona, Santiago; Iwasaki, Wataru; et al. (2022). "An Open and Continuously Updated Fern Tree of Life". Frontiers in Plant Science. 13: 909768. doi:10.3389/fpls.2022.909768. PMC 9449725. PMID 36092417.
- ^ "Tree viewer: interactive visualization of FTOL". FTOL v1.4.0 [GenBank release 253]. 2023. Retrieved 8 March 2023.
- USDA Plants Profile: Distribution in U.S.A.
- E.B.Copeland. 1947. Genera Filicum. Waltham.
- C.Michael Hogan. 2010. Fern. Encyclopedia of Earth. eds. Saikat Basu and C.Cleveland. National Council for Science and the Environment. Washington DC.
- H.A.Hyde, A.E.Wade, & S.G.Harrison. 1978. Welsh Ferns. National Museum of Wales. ISBN 0-7200-0210-9.