Vittarioideae

{{Short description|Subfamily of ferns}} {{automatic taxobox | image = Adiantum lunulatum W2 IMG_2337.jpg | image_caption = ''Adiantum lunulatum'' | taxon = Vittarioideae | authority = (C.Presl) Crabbe, Jermy & Mickel 1975 | subdivision_ranks = Genera | subdivision = See text. | synonyms = ''Adiantoideae'' {{au|(C.Presl) R.M.Tryon}} }} '''Vittarioideae''' is a subfamily of the fern family Pteridaceae, in the order Polypodiales.<ref name=PPGI/><ref name=Christenhusz-2011/> The subfamily includes the previous families '''Adiantaceae''' (adiantoids or maidenhair ferns) and '''Vittariaceae''' (vittarioids or shoestring ferns).<ref name=Smith-2006/>

==Description== The subfamily includes two distinct groups of ferns: the adiantoids, consisting of the single genus ''Adiantum'', and the vittarioids, several genera, including ''Vittaria'', which typically have highly reduced leaves, usually entire, and an epiphytic habit. The ferns historically considered as ''Adiantum'' include both petrophilic and terrestrial plants. The vittarioid ferns are primarily epiphytic in tropical regions and all have simple leaves with sori that follow the veins and lack true indusia; the sori are most often marginal with a false indusium formed from the reflexed leaf margin. The family also includes a species, ''Vittaria appalachiana'', that is highly unusual in that the sporophyte stage of the life cycle is absent. This species consists solely of photosynthetic gametophytes that reproduce asexually.{{cn|date=January 2023}}

==Taxonomy== Molecular phylogenetic analysis demonstrated that the vittarioid ferns were nested within the genus ''Adiantum'' as it was originally circumscribed, making that genus paraphyletic. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family is treated as the subfamily Vittarioideae] of the family Pteridaceae.<ref name=PPGI/>

The following diagram shows a likely phylogenetic relationship between the Vittarioideae and other subfamilies of the Pteridaceae.<ref>[http://www.pryerlab.net/publication/fichier1109.pdf Schuettpelz & Pryer (2008) "Fern phylogeny" in ''Biology and Evolution of Ferns and Lycophytes'']{{Dead link|date=July 2018 |bot=InternetArchiveBot |fix-attempted=yes }}, ed. Tom A. Ranker and Christopher H. Haufler. Cambridge University Press 2008</ref><ref name=Schu07>[http://www.pryerlab.net/publication/fichier1047.pdf Schuettpelz ''et al.'' (2007)] {{Webarchive|url=https://web.archive.org/web/20080820183832/http://www.pryerlab.net/publication/fichier1047.pdf |date=2008-08-20 }} Eric Schuettpelz, Harald Schneider, Layne Huiet, Michael D. Windham, Kathleen M. Pryer: "A molecular phylogeny of the fern family Pteridaceae: Assessing overall relationships and the affinities of previously unsampled genera." ''Molecular Phylogenetics and Evolution'' '''44''' (2007) 1172–1185</ref> {{clade|style=font-size:90%;line-height:100% |label1 = '''Pteridaceae''' |1={{clade |1=''Cryptogrammoideae'' |2={{clade |1={{clade |1=''Parkerioideae'' (syn. Ceratopteridoideae) |2=''Pteridoideae''}} |2={{clade |1=''Cheilanthoideae'' |2=''Vittarioideae''}} }} }} }}

===History=== The first suprageneric classification based on ''Vittaria'' was made by Carl Borivoj Presl in 1836, who erected the tribe '''Vittariaceae''' to contain the genera ''Vittaria'' and ''Prosaptia'', the latter now included in the grammitid ferns. He invented the new genus ''Haplopteris'' to accommodate another group of simple-leaved ferns separated from ''Pteris'', but placed it in tribe Adiantaceae instead, due to the location of its sori just behind the leaf margin.<ref>{{cite book | last=Presl | first=Carl Borivoj | authorlink=Carl Borivoj Presl | title=Tentamen Pteridologiae | year=1836 | volume=1 | publisher=Filiorum Theophili Haase | location=Prague | pages=141, 164 | url=https://www.biodiversitylibrary.org/page/32643995}}</ref>

In his 1911 treatment of the tribe, Ralph Benedict adopted a circumscription similar to modern treatments, within which he recognized the genera ''Ananthacorus'', ''Anetium'', ''Antrophyum'', ''Hecistopteris'', ''Monogramma'', ''Polytaenium'', and ''Vittaria''. He described ''Radiovittaria'' as a subgenus of ''Vittaria'', subsumed ''Scoliosorus'' within ''Polytaenium'' as doubtfully worthy of subgeneric rank, while ''Rheopteris'' had not yet been discovered.<ref>{{cite journal | last=Benedict | first=Ralph C. | title=The genera of the fern tribe Vittarieae | journal=Bulletin of the Torrey Botanical Club | volume=38 | issue=4 | year=1911 | pages=153–190 | url=https://www.biodiversitylibrary.org/page/12617283| doi=10.2307/2479298 | jstor=2479298 | hdl=2027/nnc1.cu56099444 | hdl-access=free }}</ref> ''Haplopteris'' he explicitly synonymized with ''Vittaria'' in 1914.<ref>{{cite journal | last=Benedict | first=Ralph C. | title=A revision of the genus ''Vittaria'' J. E. Smith | journal=Bulletin of the Torrey Botanical Club | volume=41 | issue=8 | year=1914 | pages=391–410 | url=https://www.biodiversitylibrary.org/page/12697291| doi=10.2307/2479721 | jstor=2479721 }}</ref>

Carl Christensen used the name "Vittarioideae" in Verdoorn's ''Manual of Pteridology'' in 1938, but did not include a description, leaving it nomenclaturally invalid. Ren-Chang Ching raised Vittariaceae to the rank of a family in 1940.<ref>{{cite journal | last=Ching | first=Ren-Chang | title=On natural classification of the family Polypodiaceae | journal=Sunyatsenia | volume=5 | year=1940 | page=232}}</ref>

The first well-sampled molecular phylogenetic study of the vittarioids was based on the chloroplast gene rbcL. In this study, it was found that the type species of ''Monogramma'' is embedded in ''Haplopteris''; the segregation of ''Vaginularia'' from ''Monogramma'' was also supported, as members of ''Vaginularia'' formed a clade sister to ''Rheopteris'' and distant from ''Monogramma'' ''sensu stricto''.<ref name="ruhfel2008">Bradley Ruhfel, Stuart Lindsay, and Charles C. Davis. 2008. "Phylogenetic Placement of ''Rheopteris'' and the Polyphyly of ''Monogramma'' (Pteridaceae s.l.): Evidence from rbcL Sequence Data". ''Systematic Botany'' '''33'''(1):37-43, {{doi|10.1600/036364408783887410}}</ref> A later molecular phylogeny, published in 2016, established the genus ''Antrophyopsis'' (formerly a subgenus of ''Antrophyum'') for three species placed in ''Scoliosorus'' but more distant from the type of that genus than ''Antrophyum''. This treatment also sank ''Anetium'' into ''Polytaenium'' and ''Monogramma'' into ''Haplopteris''.<ref>{{cite journal |last1=Schuettpelz |first1=Eric |last2=Chen |first2=Cheng-Wei |last3=Kessler |first3=Michael |last4=Pinson |first4=Jerald B. |last5=Johnson |first5=Gabriel |last6=Davila |first6=Alex |last7=Cochran |first7=Alyssa T. |last8=Huiet |first8=Layne |last9=Pryer |first9=Kathleen M. |title=A revised generic classification of vittarioid ferns (Pteridaceae) based on molecular, micromorphological, and geographic data |journal=Taxon |volume=65 |issue=4 |pages=708–722 |date=August 2016 |doi=10.12705/654.2 |bibcode=2016Taxon..65..708S |url=https://www.zora.uzh.ch/id/eprint/125839/1/Taxon_2016_708-722.pdf }}</ref> Since the name ''Monogramma'' has taxonomic priority over ''Haplopteris'', a proposal to reject ''Monogramma'' in favor of ''Haplopteris'' has been put forth to conserve the name and comparatively stable circumscription of ''Haplopteris''.<ref>{{cite journal |last1=Chen |first1=Cheng Wei |last2=Schuettpelz |first2=Eric |last3=Lindsay |first3=Stuart |last4=Middleton |first4=David J.|title=Proposal to conserve the name Haplopteris against Monogramma (Pteridaceae) |journal=Taxon |volume=65 |issue=4 |pages=884–885 |date=August 2016|doi=10.12705/654.19 |doi-access=free }}</ref>

==Genera== The following genera are recognized in the Pteridophyte Phylogeny Group classification of 2016 (PPG I):<ref name=PPGI/> * ''Adiantum'' L. * ''Ananthacorus'' Underw. & Maxon * ''Antrophyopsis'' (Benedict) Schuettp. * ''Antrophyum'' Kaulf. * ''Haplopteris'' C.Presl * ''Hecistopteris'' J.Sm. * ''Polytaenium'' Desv. * ''Radiovittaria'' (Benedict) E.H.Crane * ''Rheopteris'' Alston * ''Scoliosorus'' T.Moore * ''Vaginularia'' Fée * ''Vittaria'' Sm.

The following phylogeny for the currently recognized genera of the subfamily was presented by Schuettpelz ''et al.'':<ref name=Schu07/> {{clade|style=line-height:100% |label1 = '''Vittarioideae''' |1={{clade |1=''Adiantum'' |2={{clade |1={{clade |1=''Rheopteris'' |2=''Vaginularia''}} |2={{clade |1={{clade |1={{clade |1=''Hecistopteris'' |2=''Radiovittaria''}} |2=''Haplopteris'' (including ''Monogramma'' s.s.)}} |2={{clade |1={{clade |1=''Antrophyopsis'' |2=''Antrophyum''}} |2={{clade |1=''Polytaenium'' |2={{clade |1={{clade |1=''Scoliosorus'' |2=''Ananthacorus''}} |2=''Vittaria''}} }} }} }} }} }} }}

== References == {{Reflist|refs= <ref name=PPGI>{{citation |mode=cs1|author=PPG I |year=2016 |title=A community-derived classification for extant lycophytes and ferns |journal=Journal of Systematics and Evolution |volume=54 |issue=6 |pages=563–603 |doi=10.1111/jse.12229|doi-access=free |bibcode=2016JSyEv..54..563. }}</ref>

<ref name=Smith-2006>{{cite journal |first1=Alan R. |last1=Smith |first2=Kathleen M. |last2=Pryer |first3=Eric |last3=Schuettpelz |first4=Petra |last4=Korall |first5=Harald |last5=Schneider |first6=Paul G. |last6=Wolf |year=2006 |title=A classification for extant ferns |journal=Taxon|volume=55|issue=3|pages=705–731 | doi = 10.2307/25065646 |url=http://www.pryerlab.net/publication/fichier749.pdf|jstor=25065646 |bibcode=2006Taxon..55..705S }}</ref>

<ref name=Christenhusz-2011>{{cite journal |first1=Maarten J. M. |last1=Christenhusz |authorlink1=Maarten J. M. Christenhusz |first2=Xian-Chun |last2=Zhang |first3=Harald |last3=Schneider |year=2011 |title=A linear sequence of extant families and genera of lycophytes and ferns |journal=Phytotaxa|volume=19|pages=7–54 |url=http://www.mapress.com/phytotaxa/content/2011/f/pt00019p054.pdf|doi=10.11646/phytotaxa.19.1.2 |bibcode=2011Phytx..19....7C }}</ref> }}

Category:Pteridaceae Category:Plant subfamilies