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{{Short description|Subfamily of ferns}}
{{Taxobox
{{automatic taxobox
| name = Vittarioideae
| image = Adiantum lunulatum W2 IMG_2337.jpg
| regnum = [[Plant]]ae
| divisio = [[Pteridophyta]]
| image_caption = ''[[Adiantum lunulatum]]''
| taxon = Vittarioideae
| classis = [[Polypodiopsida]]
| authority = (C.Presl) Crabbe, Jermy & Mickel 1975
| ordo = [[Polypodiales]]
| familia = [[Pteridaceae]]
| subfamilia = '''Vittarioideae'''
| subfamilia_authority = (C.Presl) Crabbe, Jermy & Mickel 1975
| subdivision_ranks = Genera
| subdivision_ranks = Genera
| subdivision = {{species list
| subdivision = [[#Genera|See text]].
| synonyms = ''Adiantoideae'' {{au|(C.Presl) R.M.Tryon}}
|[[Adiantum]]|
|[[Ananthacorus]]|
|[[Anetium]]|
|[[Antrophyum]]|
|[[Haplopteris]]|
|[[Hecistopteris]]|
|[[Monogramma]]|
|[[Polytaenium]]|
|[[Radiovittaria]]|
|[[Rheopteris]]|
|[[Scoliosorus]]|
|[[Vittaria]]|<ref name=Christenhusz-2011/>}}
| synonyms = ''[[Adiantoideae]]'' {{au|(C.Presl) R.M.Tryon 1986}}
}}
}}
'''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==
'''Vittarioideae''' is a subfamily of the [[Pteridaceae]] family of ferns,<ref name=Christenhusz-2011/> in the order [[Polypodiales]], class [[Polypodiopsida]]. This subfamily includes the previous families [[Adiantaceae]] (adiantoids or maidenhair ferns) and [[Vittariaceae]] (vittarioids or shoestring ferns).<ref name=Smith-2006/>
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 [[epiphyte|epiphytic]] in tropical regions and all have simple leaves with [[sorus|sori]] that follow the veins and lack true [[indusium|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 [[photosynthesis|photosynthetic]] [[gametophyte]]s that reproduce asexually.{{cn|date=January 2023}}


==Taxonomy==
The following diagram, shows a likely phylogenic 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''], ed. Tom A. Ranker and Christopher H. Haufler. Cambridge University Press 2008</ref><ref>[http://www.pryerlab.net/publication/fichier1047.pdf Schuettpelz ''et al.'' (2007)] 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>
[[Molecular phylogenetics|Molecular phylogenetic]] analysis demonstrated that the vittarioid ferns were nested within the genus ''[[Adiantum]]'' as it was originally [[Circumscription (taxonomy)|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%
{{clade|style=font-size:90%;line-height:100%
|label1 = '''[[Pteridaceae]]'''
|label1 = '''[[Pteridaceae]]'''
|1={{clade|
|1={{clade
|1=''[[Cryptogrammoideae]]''
|1=''[[Cryptogrammoideae]]''
|2={{clade
|2={{clade
|1={{clade
|1={{clade
|1=''[[Ceratopteridoideae]]''
|1=''[[Parkerioideae]]'' (syn. Ceratopteridoideae)
|2=''[[Pteridoideae]]''}}
|2=''[[Pteridoideae]]''}}
|2={{clade
|2={{clade
|1=''[[Cheilanthoideae]]''
|1=''[[Cheilanthoideae]]''
|2=''[[Vittarioideae]]''}}
|2=''Vittarioideae''}}
}}
}}
}}
}}
}}
}}
==References==
{{reflist|refs=
<ref name=Smith-2006>{{cite journal
|author=Alan R. Smith, Kathleen M. Pryer, Eric Schuettpelz, Petra Korall, Harald Schneider & Paul G. 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}}</ref>


===History===
<ref name=Christenhusz-2011>{{cite journal
The first suprageneric classification based on ''Vittaria'' was made by [[Carl Borivoj Presl]] in 1836, who erected the [[tribe (biology)|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 | publisher=Filiorum Theophili Haase | location=Prague | pages=141, 164 | url=https://www.biodiversitylibrary.org/page/32643995}}</ref>
|author=Maarten J. M. Christenhusz, Xian-Chun Zhang & Harald Schneider

|year=2011
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 (plant)|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>
|title=A linear sequence of extant families and genera of lycophytes and ferns

|journal=[[Phytotaxa]]|volume=19|pages=7–54
[[Carl Christensen (botanist)|Carl Christensen]] used the name "Vittarioideae" in [[Inez Clare Verdoorn|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> <!-- what was his circumscription? -->
|url=http://www.mapress.com/phytotaxa/content/2011/f/pt00019p054.pdf}}</ref>

The first well-sampled [[molecular phylogenetic]] [[Research|study]] of the vittarioids was based on the [[chloroplast]] [[gene]] [[RuBisCO large subunit|rbcL]]. In this study, it was found that the [[type species]] of ''Monogramma'' is [[Paraphyly|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 |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]]'' <small>L.</small>
* ''[[Ananthacorus]]'' <small>Underw. & Maxon</small>
* ''[[Antrophyopsis]]'' <small>(Benedict) Schuettp.</small>
* ''[[Antrophyum]]'' <small>Kaulf.</small>
* ''[[Haplopteris]]'' <small>C.Presl</small>
* ''[[Hecistopteris]]'' <small>J.Sm.</small>
* ''[[Polytaenium]]'' <small>Desv.</small>
* ''[[Radiovittaria]]'' <small>(Benedict) E.H.Crane</small>
* ''[[Rheopteris]]'' <small>Alston</small>
* ''[[Scoliosorus]]'' <small>T.Moore</small>
* ''[[Vaginularia]]'' <small>Fée</small>
* ''[[Vittaria]]'' <small>Sm.</small>

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 (plant)|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 }}</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 }}</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 }}</ref>
}}
}}


{{Taxonbar|from1=Q5873935|from2=Q19387804|from3=Q3735428}}
[[Category:Pteridaceae| ]]
{{fern-stub}}


[[Category:Pteridaceae]]
[[es:Vittarioideae]]
[[Category:Plant subfamilies]]

Latest revision as of 22:41, 14 August 2023

Vittarioideae
Adiantum lunulatum
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Division: Polypodiophyta
Class: Polypodiopsida
Order: Polypodiales
Family: Pteridaceae
Subfamily: Vittarioideae
(C.Presl) Crabbe, Jermy & Mickel 1975
Genera

See text.

Synonyms

Adiantoideae (C.Presl) R.M.Tryon

Vittarioideae is a subfamily of the fern family Pteridaceae, in the order Polypodiales.[1][2] The subfamily includes the previous families Adiantaceae (adiantoids or maidenhair ferns) and Vittariaceae (vittarioids or shoestring ferns).[3]

Description[edit]

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.[citation needed]

Taxonomy[edit]

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.[1]

The following diagram shows a likely phylogenetic relationship between the Vittarioideae and other subfamilies of the Pteridaceae.[4][5]

Pteridaceae

Cryptogrammoideae

Parkerioideae (syn. Ceratopteridoideae)

Pteridoideae

Cheilanthoideae

Vittarioideae

History[edit]

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.[6]

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.[7] Haplopteris he explicitly synonymized with Vittaria in 1914.[8]

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.[9]

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.[10] 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.[11] 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.[12]

Genera[edit]

The following genera are recognized in the Pteridophyte Phylogeny Group classification of 2016 (PPG I):[1]

The following phylogeny for the currently recognized genera of the subfamily was presented by Schuettpelz et al.:[5]

Vittarioideae

References[edit]

  1. ^ a b c PPG I (2016). "A community-derived classification for extant lycophytes and ferns". Journal of Systematics and Evolution. 54 (6): 563–603. doi:10.1111/jse.12229.
  2. ^ Christenhusz, Maarten J. M.; Zhang, Xian-Chun; Schneider, Harald (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.
  3. ^ Smith, Alan R.; Pryer, Kathleen M.; Schuettpelz, Eric; Korall, Petra; Schneider, Harald; Wolf, Paul G. (2006). "A classification for extant ferns" (PDF). Taxon. 55 (3): 705–731. doi:10.2307/25065646. JSTOR 25065646.
  4. ^ Schuettpelz & Pryer (2008) "Fern phylogeny" in Biology and Evolution of Ferns and Lycophytes[permanent dead link], ed. Tom A. Ranker and Christopher H. Haufler. Cambridge University Press 2008
  5. ^ a b Schuettpelz et al. (2007) Archived 2008-08-20 at the Wayback Machine 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
  6. ^ Presl, Carl Borivoj (1836). Tentamen Pteridologiae. Prague: Filiorum Theophili Haase. pp. 141, 164.
  7. ^ Benedict, Ralph C. (1911). "The genera of the fern tribe Vittarieae". Bulletin of the Torrey Botanical Club. 38 (4): 153–190. doi:10.2307/2479298. hdl:2027/nnc1.cu56099444. JSTOR 2479298.
  8. ^ Benedict, Ralph C. (1914). "A revision of the genus Vittaria J. E. Smith". Bulletin of the Torrey Botanical Club. 41 (8): 391–410. doi:10.2307/2479721. JSTOR 2479721.
  9. ^ Ching, Ren-Chang (1940). "On natural classification of the family Polypodiaceae". Sunyatsenia. 5: 232.
  10. ^ 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
  11. ^ Schuettpelz, Eric; Chen, Cheng-Wei; Kessler, Michael; Pinson, Jerald B.; Johnson, Gabriel; Davila, Alex; Cochran, Alyssa T.; Huiet, Layne; Pryer, Kathleen M. (August 2016). "A revised generic classification of vittarioid ferns (Pteridaceae) based on molecular, micromorphological, and geographic data" (PDF). Taxon. 65 (4): 708–722. doi:10.12705/654.2.
  12. ^ Chen, Cheng Wei; Schuettpelz, Eric; Lindsay, Stuart; Middleton, David J. (August 2016). "Proposal to conserve the name Haplopteris against Monogramma (Pteridaceae)". Taxon. 65 (4): 884–885. doi:10.12705/654.19.
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