Jump to content

Thelypteridaceae: Difference between revisions

From Wikipedia, the free encyclopedia
Browse history interactively
← Previous edit
Content deleted Content added
VisualWikitext
→Classification: subfamily names
 
(27 intermediate revisions by 14 users not shown)
Line 1: Line 1:
{{Short description|Family of ferns}}
{{automatic taxobox
{{Automatic taxobox
| name = Marsh fern family
| name = Marsh fern family
| oldest_fossil = Cenomanian
| image =Starr_041221-1892_Pneumatopteris_sandwicensis.jpg
| image = Starr_041221-1892_Pneumatopteris_sandwicensis.jpg
| image_caption = ''[[Cyclosorus]] cyatheoides''
| image_caption = ''[[Cyclosorus]] cyatheoides''
| taxon = Thelypteridaceae
| taxon = Thelypteridaceae
| authority = [[Ren Chang Ching|Ching]] ex Pic. Serm. 1970<ref>[http://www.tropicos.org/Name/42000412 Thelypteridaceae Ching ex Pic. Serm.] Tropicos.org. Missouri Botanical Garden. 24 Jan 2012</ref>
| authority = [[Ren Chang Ching|Ching]] ex Pic.Serm.<ref>[http://www.tropicos.org/Name/42000412 Thelypteridaceae Ching ex Pic. Serm.] Tropicos.org. Missouri Botanical Garden. 24 Jan 2012</ref>
| subdivision_ranks = Genera
| subdivision_ranks = Genera
| subdivision = {{species list
| subdivision = See text.
|[[Cyclosorus]]|Link 1833
|[[Macrothelypteris]]|(H.Itô) Ching 1963
|[[Phegopteris]]|(C.Presl) Fée 1852
|[[Pseudophegopteris]]|Ching 1963
|[[Thelypteris]]|Ching 1963}}
}}
}}


'''Thelypteridaceae''' is a family of about 900<ref name=FNA/> species of [[fern]]s in the order [[Polypodiales]].<ref name=PPGI/><ref name=Christenhusz-2011/><ref name=Smith-2006/> In the [[Pteridophyte Phylogeny Group]] classification of 2016 (PPG I), it is placed in the suborder [[Aspleniineae]].<ref name=PPGI/> Alternatively, the family may be submerged in a very broadly defined family Aspleniaceae as the subfamily '''Thelypteridoideae'''.<ref name=ChriChas14/>
'''Thelypteridaceae''' is a family of about 900<ref name=FNA/> species of [[fern]]s in the order [[Polypodiales]].<ref name=PPGI/><ref name=Christenhusz-2011/><ref name=Smith-2006/> In the [[Pteridophyte Phylogeny Group]] classification of 2016 (PPG I), it is placed in the suborder [[Aspleniineae]].<ref name=PPGI/> Alternatively, the family may be submerged in a very broadly defined family Aspleniaceae as the subfamily '''Thelypteridoideae'''.<ref name=ChriChas14/>


The ferns are [[Terrestrial plant|terrestrial]], with the exception of a few which are [[lithophyte]]s (grow on rocks).<ref name=Smith-2006/> The bulk of the species are tropical, although there are a number of temperate species.<ref name=Smith-2006/>
The ferns are [[Terrestrial plant|terrestrial]], with the exception of a few which are [[lithophyte]]s (grow on rocks).<ref name=Smith-2006/> The bulk of the species are tropical, although there are a number of temperate species.<ref name=Smith-2006/>


These ferns typically have creeping [[rhizome]]s. The [[frond]]s are simply [[pinnate]] to pinnate-[[pinnatifid]]. There is either no frond [[Sexual dimorphism|dimorphism]] or only mild dimorphism, either open [[Leaf#Venation .28arrangement of the veins.29|venation]] or very simple [[anastomosing]]. The [[sorus|sori]] are mostly reniform in shape and have indusia, except for the ''Phegopteris'' group.
These ferns typically have creeping [[rhizome]]s. The [[frond]]s are simply [[pinnate]] to pinnate-[[pinnatifid]]. There is either no frond [[Sexual dimorphism|dimorphism]] or only mild dimorphism, either open [[Leaf#Venation .28arrangement of the veins.29|venation]] or very simple [[anastomosing]]. The [[sorus|sori]] are mostly reniform in shape and have indusia, except for the ''Phegopteris'' group.


== Classification ==
== Classification ==
At one time,{{specify|date=March 2011}} all thelypterioid ferns were included in the genus ''[[Dryopteris]]'' because of the sorus shape. However, there are a great many differences between the groups, and these plants are now segregated in their own family.<ref name=FNA/>
During the early and mid 1900's<ref name="ferns2001">{{cite journal |last1=Collinson |first1=M. E. |title=Cainozoic ferns and their distribution |journal=Brittonia |date=2001 |volume=53 |issue=2 |pages=173–235 |doi=10.1007/BF02812700 |jstor=2666600|s2cid=19984401 }}</ref> all thelypterioid ferns were included in the genus ''[[Dryopteris]]'' because of the sorus shape. However, there are a great many differences between the groups, and these plants are now segregated in their own family.<ref name=FNA/><ref name=PPGI/> [[Genetics|Genetic]] evidence shows that the family is clearly [[Monophyly|monophyletic]].<ref name=PPGI/>


===Phylogenetic relationships===
Some researchers include the entire family Thelypteridaceae in the genus ''[[Thelypteris]]''; others divide the family into as many as 30 genera. An intermediate position is to place the bulk in ''Thelypteris'' (which can then be divided into subgenera and sections corresponding to the genera of other authors) but to separate out ''[[Phegopteris]]'' and ''[[Macrothelypteris]]''.<ref name=FNA/> Another choice is to divide the family into a half a dozen or so genera.<ref name=Smith-2006/>
The following cladogram for the suborder [[Aspleniineae]] (as eupolypods II), based on Lehtonen (2011),<ref name=Lehtonen-2011/> and Rothfels & al. (2012),<ref name=Rothfels-2012/> shows a likely [[phylogenetics|phylogenetic]] relationship between the Thelypteridaceae and the other families of the clade.
{{eupolypods II cladogram}}


===Genera===
This family includes several complexes of species that are difficult to distinguish, and seem to represent a remarkable evolutionary radiation.
The family can be divided into two major clades, which the [[Pteridophyte Phylogeny Group]] classification of 2016 (PPG I) recognizes as two subfamilies, Phegopteridoideae and Thelypteroideae.<ref name=PPGI/> Their division into genera has been described as "highly controversial and fluctuating".<ref name=CFLW_Th/> The family includes several complexes of species that are difficult to distinguish, and seem to represent a remarkable evolutionary radiation. Some researchers include the entire family Thelypteridaceae in the genus ''[[Thelypteris]]'';{{citation needed|date=January 2020}} others divide the family into as many as 30 genera.<ref name=CFLW_Th/> An intermediate position is to place the bulk in ''Thelypteris'' (which can then be divided into subgenera and sections corresponding to the genera of other authors) but to separate out ''[[Phegopteris]]'' and ''[[Macrothelypteris]]''.<ref name=FNA/> Another choice is to divide the family into a half a dozen or so genera.<ref name=Smith-2006/>


{{cladogram|title=|Phylogeny of Thelypteriaceae<ref>{{cite journal |last1=Nitta |first1=Joel H. |last2=Schuettpelz |first2=Eric |last3=Ramírez-Barahona |first3=Santiago |last4=Iwasaki |first4=Wataru |display-authors=et al. |year=2022 |title=An Open and Continuously Updated Fern Tree of Life |journal=Frontiers in Plant Science |volume=13 |issue= |page= 909768| doi=10.3389/fpls.2022.909768 |pmid= 36092417|pmc= 9449725|bibcode= |doi-access=free}}</ref><ref>{{cite web|last1= |first1= |last2= |display-authors=et al. |year=2022 |title=Tree viewer: interactive visualization of FTOL |url=https://fernphy.github.io/viewer.html |version=FTOL v1.3.0 |access-date=12 December 2022}}</ref>
Recent [[Genetics|genetic]] evidence shows that the family is clearly [[cladistics|monophyletic]]. The same genetic studies show that the traditional genus ''Phegopteris'' is, in fact, a clearly segregate group that diverges at the very base of the [[family tree]]. In fact, this particular [[clade]] also includes the genera ''[[Pseudophegopteris]]'' and ''[[Macrothelypteris]]'', which are here included in the larger genus ''[[Phegopteris]]''. Similarly, ''Coryphopteris'' is included here in the genus ''[[Metathelypteris]]'', ''Parathelypteris'' is included in ''[[Amauropelta]]'', and a large group of genera are included in ''[[Cyclosorus]]''.
{{Clade | style=font-size:90%;line-height:80%;width:500px
|1={{clade
|label1=Phegopteridoideae
|1={{clade
|1=''[[Macrothelypteris]]''
|2={{clade
|1=''[[Phegopteris]]''
|2=''[[Pseudophegopteris]]''
}}
}}
|label2=Thelypteridoideae
|2={{clade
|1={{clade
|label1=Thelypterideae
|1=''[[Thelypteris]]''
}}
|2={{clade
|1={{clade
|label1=Amauropelteae
|1={{clade
|1=''[[Coryphopteris]]''
|2={{clade
|1=''[[Metathelypteris]]''
|2=''[[Amauropelta]]''
}}
}}
}}
|2={{clade
|1=''[[Oreopteris]]''
|2={{clade
|1=''[[Steiropteris]]''
|2={{clade
|label1=Leptogrammeae
|1={{clade
|1=''[[Cyclogramma]]''
|2={{clade
|1=''[[Stegnogramma]]''
|2=''[[Leptogramma]]''
}}
}}
|label2=Meniscieae
|2={{clade
|1=''[[Steiropteris]]'' species-group 2
|2={{clade
|1={{clade
|label1=Goniopteridinae
|1=''[[Goniopteris]]''
}}
|2={{clade
|label1=Menisciinae
|1={{clade
|1={{clade
|1=''[[Cyclosorus]]''
|2={{clade
|1=''[[Ampelopteris]]''
|2=''[[Mesophlebion]]''
}}
}}
|2=''[[Meniscium]]''
}}
|label2=Pseudocyclosorinae
|2={{clade
|1=''[[Pelazoneuron]]''
|2={{clade
|1=''[[Pakau]]''
|2={{clade
|1=''Pelazoneuron'' species-group 2
|2={{clade
|1=''[[Glaphyropteridopsis]]''
|2={{clade
|1={{clade
|1=''[[Chingia]]''
|2={{clade
|1=''[[Plesioneuron]]''
|2=''[[Menisciopsis]]''
}}
}}
|2={{clade
|1=''[[Grypothrix]]''
|2={{clade
|1=''[[Mesopteris]]''
|2={{clade
|1=''[[Plesioneuron archboldiae]]''
|2={{clade
|1=''[[Strophocaulon]]''
|2={{clade
|1={{clade
|1={{clade
|1=''[[Pneumatopteris]]''
|2=''[[Reholttumia]]''
}}
|2={{clade
|1=''[[Pronephrium]]''
|2=''[[Sphaerostephanos]]''
}}
}}
|2={{clade
|1={{clade
|1=''[[Trigonospora]]''
|2=''[[Pseudocyclosorus]]''
}}
|2={{clade
|1={{clade
|1=''[[Amblovenatum]]''
|2={{clade
|1=''[[Sphaerostephanos elatus]]''
|2=''[[Abacopteris]]''
}}
}}
|2=''[[Christella]]''
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}


The Pteridophyte Phylogeny Group classification of 2016 (PPG I) accepts 30 genera:<ref name=PPGI/>
The most recent molecular phylogenetic study divides the family into two major clades:
*Subfamily Phegopteridoideae <small>Salino, A.R.Sm. & T.E.Almeid</small>
*The phegopteroids, subfamily Phegopteridoideae, comprising the genera ''[[Macrothelypteris]]'', ''[[Phegopteris]]'' and ''[[Pseudophegopteris]]''
**''[[Macrothelypteris]]'' <small>(H.Ito) Ching</small>
*The thelypteroids, subfamily Thelypteroideae, which include ''[[Thelypteris]]'' s.s., the amauropeltoid clade (''[[Amauropelta]]'', ''[[Coryphopteris]]'', ''[[Metathelypteris]]'', and ''[[Parathelypteris]]''), ''[[Oreopteris]]'', and the cyclosoroid clade (the christelloids, ''[[Ampelopteris]]'', ''[[Cyclogramma]]'', ''[[Cyclosorus]]'' s.s., ''[[Goniopteris]]'', ''[[Meniscium]]'', ''[[Mesophlebion]]'', ''[[Stegnogramma]]'' s.l., and ''[[Steiropteris]]''). The christelloids comprise ''[[Amphineuron]]'', ''[[Chingia]]'', ''[[Christella]]'', ''[[Glaphyropteridopsis]]'', ''[[Mesopteris]]'', ''[[Plesioneuron]]'', ''[[Nannothelypteris]]'', ''[[Pneumatopteris]]'', ''[[Pronephrium]]'', ''[[Pseudocyclosorus]]'', ''[[Sphaerostephanos]]'', ''[[Trigonospora]]'', and part of ''Cyclosorus'' s.l.; current generic boundaries in the christelloids will require re-circumscription after further phylogenetic study.
**''[[Phegopteris]]'' <small>(C.Presl) Fée</small>
**''[[Pseudophegopteris]]'' <small>Ching</small><ref name=CFLW_Ph/><ref name=POWO_30000947-2/>
*Subfamily Thelypteridoideae <small>C.F.Reed</small>
**''[[Steiropteris]]'' <small>(C.Chr.) Pic.Serm.</small>
**Tribe Thelypterideae <small>Ching 1963</small>
***''[[Thelypteris]]'' <small>Schmidel</small>
**Tribe Amauropelteae <small>Wei, Liu & Liu</small>
***''[[Amauropelta]]'' <small>Kunze</small>
***''[[Coryphopteris]]'' <small>Holttum</small>
***''[[Metathelypteris]]'' <small>(H.Ito) Ching</small>
**Tribe Oreopterideae <small>Wei, Liu & Liu</small>
***''[[Oreopteris]]'' <small>Holub</small>
**Tribe Leptogrammeae <small>Todaro 1866</small>
***''[[Cyclogramma]]'' <small>Tagawa</small>
***''[[Leptogramma]]'' <small>Smith</small>
***''[[Stegnogramma]]'' <small>Blume</small>
**Tribe Meniscieae <small>Fée 1850-1852 ex Pfeiffer 1873</small>
***Subtribe Goniopteridinae <small>Ching 1963</small>
****''[[Goniopteris]]'' <small>C.Presl</small>
***Subtribe Menisciinae <small>Payer 1850</small>
****''[[Ampelopteris]]'' <small>Kunze</small>
****''[[Cyclosorus]]'' <small>Link</small>
****''[[Meniscium]]'' <small>Schreb.</small>
****''[[Mesophlebion]]'' <small>Holttum</small>
***Subtribe Pseudocyclosorinae <small>Ching 1963</small>
****''[[Abacopteris]]'' <small>Fée</small>
****''[[Amblovenatum]]'' <small>J.P.Roux</small>
****''[[Chingia]]'' <small>Holttum</small>
****''[[Christella]]'' <small>H.Lév.</small>
****''[[Glaphyropteridopsis]]'' <small>Ching</small>
****''[[Grypothrix]]'' <small>(Holttum 1971) Fawcett & Smith</small>
****''[[Menisciopsis]]'' <small>(Holttum 1982) Fawcett & Smith</small>
****''[[Menisorus]]'' <small>Alston</small>
****''[[Mesopteris]]'' <small>Ching</small>
****''[[Pakau]]'' <small>Fawcett & Smith</small>
****''[[Parathelypteris]]'' <small>(H.Ito) Ching</small>
****''[[Pelazoneuron]]'' <small>(Holttum 1974) Smith & Fawcett</small>
****''[[Plesioneuron]]'' <small>(Holttum) Holttum</small>
****''[[Pneumatopteris]]'' <small>Nakai</small>
****''[[Pronephrium]]'' <small>C.Presl</small> [''[[Nannothelypteris]]'' <small>Holttum</small>]
****''[[Pseudocyclosorus]]'' <small>Ching</small>
****''[[Reholttumia]]'' <small>Fawcett & Smith</small>
****''[[Sphaerostephanos]]'' <small>J.Sm.</small>
****''[[Strophocaulon]]'' <small>Fawcett & Smith</small>
****''[[Trigonospora]]'' <small>Holttum</small>
Some species of ''Pronephrium'' were split off into other genera in 2021 as a result of a [[Phylogenetics|phylogenetic]] study of the family Thelypteridaceae:<ref name=FawcSmit21>{{Citation |last1=Fawcett |first1=S. |last2=Smith |first2=A.R. |date=2021 |title=A Generic Classification of the Thelypteridaceae |series=Sida, Botanical Miscellany 59 |publication-place=Fort Worth |publisher=Botanical Research Institute of Texas Press |name-list-style=amp }}</ref><ref name=FawcSmitSundBurl21>{{Citation |last1=Fawcett |first1=Susan |last2=Smith |first2=Alan R. |last3=Sundue |first3=Michael |last4=Burleigh |first4=J. Gordon |last5=Sessa |first5=Emily B. |last6=Kuo |first6=Li-Yaung |last7=Chen |first7=Cheng-Wei |last8=Testo |first8=Weston L. |last9=Kessler |first9=Michael |last10=Barrington |first10=David S. |date=2021 |title=A Global Phylogenomic Study of the Thelypteridaceae |journal=Systematic Botany |volume=46 |issue=4 |pages=891–915 |url=https://sessalab.org/wp-content/uploads/2022/01/Thelyp2021.pdf |access-date=2022-06-04 |doi=10.1600/036364421X16370109698650 |s2cid=245512861 |name-list-style=amp }}</ref>
*''[[Abacopteris]]'' <small>Fée</small>
*''[[Grypothrix]]'' <small>(Holttum) S.E.Fawc. & A.R.Sm.</small>
*''[[Menisciopsis]]'' <small>(Holttum) S.E.Fawc. & A.R.Sm</small>


The extinct genus ''[[Holttumopteris]]'' from the [[Cenomanian]] aged [[Burmese amber]] has been suggested to have affinities with the family, but several important diagnostic characters are not preserved.<ref>{{Cite journal|last1=Regalado|first1=Ledis|last2=Schmidt|first2=Alexander R.|last3=Krings|first3=Michael|last4=Bechteler|first4=Julia|last5=Schneider|first5=Harald|last6=Heinrichs|first6=Jochen|date=January 2018|title=Fossil evidence of eupolypod ferns in the mid-Cretaceous of Myanmar|url=http://link.springer.com/10.1007/s00606-017-1439-2|journal=Plant Systematics and Evolution|language=en|volume=304|issue=1|pages=1–13|doi=10.1007/s00606-017-1439-2|s2cid=21617872|issn=0378-2697}}</ref>
===Phylogenetic relationships===
The following cladogram for the suborder [[Aspleniineae]] (as eupolypods II), based on Lehtonen (2011),<ref name=Lehtonen-2011/> and Rothfels & al. (2012),<ref name=Rothfels-2012/> shows a likely [[phylogenetics|phylogenetic]] relationship between the Thelypteridaceae and the other families of the clade.
{{eupolypods II cladogram}}


==References==
==References==
{{Reflist|refs=
{{Reflist|refs=
<ref name=ChriChas14>{{Cite journal |last1=Christenhusz |first1=Maarten J.M. |last2=Chase |first2=Mark W. |date=2014 |title=Trends and concepts in fern classification |journal=Annals of Botany |volume=113 |issue=9 |pages=571–594 |doi=10.1093/aob/mct299 |pmid=24532607 |pmc=3936591 |lastauthoramp=yes }}</ref>
<ref name=CFLW_Th>{{Citation |mode=cs1 |last1=Hassler |first1=Michael |last2=Schmitt |first2=Bernd |date=January 2020 |contribution=Thelypteridaceae |title=Checklist of Ferns and Lycophytes of the World |at=8.20 |url=https://worldplants.webarchiv.kit.edu/ferns/ |access-date=2020-01-11 |name-list-style=amp }}</ref>


<ref name=CFLW_Ph>{{Citation |mode=cs1 |last1=Hassler |first1=Michael |last2=Schmitt |first2=Bernd |date=January 2020 |contribution=Phegopteris |title=Checklist of Ferns and Lycophytes of the World |at=8.20 |url=https://worldplants.webarchiv.kit.edu/ferns/ |access-date=2020-01-10 |name-list-style=amp }}</ref>
<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
|url=http://www.pryerlab.net/publication/fichier749.pdf
|doi=10.2307/25065646}}</ref>


<ref name=Christenhusz-2011>{{cite journal
<ref name=Christenhusz-2011>{{cite journal
|author=Maarten J. M. Christenhusz, Xian-Chun Zhang & Harald Schneider
|author=Maarten J. M. Christenhusz
|author2=Xian-Chun Zhang
|author3=Harald Schneider
|name-list-style=amp
|year=2011
|year=2011
|title=A linear sequence of extant families and genera of lycophytes and ferns
|title=A linear sequence of extant families and genera of lycophytes and ferns
|journal=[[Phytotaxa]]|volume=19|pages=7–54
|journal=[[Phytotaxa]]|volume=19|pages=7–54
|doi=10.11646/phytotaxa.19.1.2
|url=http://www.mapress.com/phytotaxa/content/2011/f/pt00019p054.pdf}}</ref>
|url=http://www.mapress.com/phytotaxa/content/2011/f/pt00019p054.pdf}}</ref>

<ref name=ChriChas14>{{Cite journal |last1=Christenhusz |first1=Maarten J.M. |last2=Chase |first2=Mark W. |date=2014 |title=Trends and concepts in fern classification |journal=Annals of Botany |volume=113 |issue=9 |pages=571–594 |doi=10.1093/aob/mct299 |pmid=24532607 |pmc=3936591 |name-list-style=amp }}</ref>

<ref name=FNA>{{cite web
|url = http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=10887
|work = Flora of North America
|title = Thelypteridaceae}}</ref>


<ref name=Lehtonen-2011>{{cite journal
<ref name=Lehtonen-2011>{{cite journal
Line 60: Line 245:
|year=2011
|year=2011
|title=Towards Resolving the Complete Fern Tree of Life
|title=Towards Resolving the Complete Fern Tree of Life
|journal=PLoS ONE |volume=6 |issue=10 |pages=e24851
|journal=PLOS ONE |volume=6 |issue=10 |pages=e24851
|url=http://www.sci.utu.fi/sivustot/amazon/publications/articles/pdf_pub/Lehtonen_2011_PLoS1.pdf |doi=10.1371/journal.pone.0024851 |pmid=22022365 |pmc=3192703}}</ref>
|url=http://www.sci.utu.fi/sivustot/amazon/publications/articles/pdf_pub/Lehtonen_2011_PLoS1.pdf |doi=10.1371/journal.pone.0024851 |pmid=22022365 |pmc=3192703|bibcode=2011PLoSO...624851L
|doi-access=free
}}</ref>


<ref name=POWO_30000947-2>{{Citation |mode=cs1 |title=''Phegopteris'' (C.Presl) Fée |work=Plants of the World Online |publisher=Royal Botanic Gardens, Kew|url=https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:30000947-2 |access-date=2020-01-10 }}</ref>
<ref name=PPGI>{{citation |mode=cs1|author=PPG I |title=A community-derived classification for extant lycophytes and ferns |year=2016 |journal=Journal of Systematics and Evolution |volume=54 |issue=6 |pages=563–603 |doi=10.1111/jse.12229}}</ref>

<ref name=PPGI>{{citation |mode=cs1|author=PPG I |title=A community-derived classification for extant lycophytes and ferns |year=2016 |journal=Journal of Systematics and Evolution |volume=54 |issue=6 |pages=563–603 |doi=10.1111/jse.12229|s2cid=39980610 |doi-access=free }}</ref>


<ref name=Rothfels-2012>{{cite journal
<ref name=Rothfels-2012>{{cite journal
|author1=Carl J. Rothfels |author2=Anders Larsson |author3=Li-Yaung Kuo |author4=Petra Korall |author5=Wen- Liang Chiou |author6=Kathleen M. Pryer |year=2012
|author1=Carl J. Rothfels |author2=Anders Larsson |author3=Li-Yaung Kuo |author4=Petra Korall |author5=Wen- Liang Chiou |author6=Kathleen M. Pryer |year=2012
|title=Overcoming Deep Roots, Fast Rates, and Short Internodes to Resolve the Ancient Rapid Radiation of Eupolypod II Ferns
|title=Overcoming Deep Roots, Fast Rates, and Short Internodes to Resolve the Ancient Rapid Radiation of Eupolypod II Ferns
|journal=Systematic Biology |volume=61 |issue=1 |pages=70
|journal=Systematic Biology |volume=61 |issue=1 |pages=490–509
|url=http://sysbio.oxfordjournals.org/content/early/2012/01/04/sysbio.sys001.short?rss=1|doi=10.1093/sysbio/sys001 |pmid=22223449|arxiv=cond-mat/0606434 }}</ref>
|url=http://sysbio.oxfordjournals.org/content/early/2012/01/04/sysbio.sys001.short?rss=1|doi=10.1093/sysbio/sys001 |pmid=22223449|arxiv=cond-mat/0606434 }}</ref>


<ref name=FNA>{{cite web
<ref name=Smith-2006>{{cite journal
|author=Alan R. Smith
|url = http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=10887
|author2=Kathleen M. Pryer
|work = Flora of North America
|author3=Eric Schuettpelz
|title = Thelypteridaceae}}</ref>
|author4=Petra Korall
|author5=Harald Schneider
|author6=Paul G. Wolf
|name-list-style=amp
|year=2006
|title=A classification for extant ferns
|journal=Taxon|volume=55|issue=3|pages=705–731
|url=http://www.pryerlab.net/publication/fichier749.pdf
|doi=10.2307/25065646|jstor=25065646
}}</ref>
}}
}}


{{Plant classification}}
{{Taxonbar|from=Q1496479}}
{{Taxonbar|from=Q1496479}}
{{Authority control}}


[[Category:Fern families]]
[[Category:Thelypteridaceae| ]]
[[Category:Thelypteridaceae| ]]
[[Category:Fern families]]

Latest revision as of 18:12, 25 October 2023

Marsh fern family
Temporal range: Cenomanian–Recent
Cyclosorus cyatheoides
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Division: Polypodiophyta
Class: Polypodiopsida
Order: Polypodiales
Suborder: Aspleniineae
Family: Thelypteridaceae
Ching ex Pic.Serm.[1]
Genera

See text.

Thelypteridaceae is a family of about 900[2] species of ferns in the order Polypodiales.[3][4][5] In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), it is placed in the suborder Aspleniineae.[3] Alternatively, the family may be submerged in a very broadly defined family Aspleniaceae as the subfamily Thelypteridoideae.[6]

The ferns are terrestrial, with the exception of a few which are lithophytes (grow on rocks).[5] The bulk of the species are tropical, although there are a number of temperate species.[5]

These ferns typically have creeping rhizomes. The fronds are simply pinnate to pinnate-pinnatifid. There is either no frond dimorphism or only mild dimorphism, either open venation or very simple anastomosing. The sori are mostly reniform in shape and have indusia, except for the Phegopteris group.

Classification

[edit]

During the early and mid 1900's[7] all thelypterioid ferns were included in the genus Dryopteris because of the sorus shape. However, there are a great many differences between the groups, and these plants are now segregated in their own family.[2][3] Genetic evidence shows that the family is clearly monophyletic.[3]

Phylogenetic relationships

[edit]

The following cladogram for the suborder Aspleniineae (as eupolypods II), based on Lehtonen (2011),[8] and Rothfels & al. (2012),[9] shows a likely phylogenetic relationship between the Thelypteridaceae and the other families of the clade.

Aspleniineae (eupolypods II)

Genera

[edit]

The family can be divided into two major clades, which the Pteridophyte Phylogeny Group classification of 2016 (PPG I) recognizes as two subfamilies, Phegopteridoideae and Thelypteroideae.[3] Their division into genera has been described as "highly controversial and fluctuating".[10] The family includes several complexes of species that are difficult to distinguish, and seem to represent a remarkable evolutionary radiation. Some researchers include the entire family Thelypteridaceae in the genus Thelypteris;[citation needed] others divide the family into as many as 30 genera.[10] An intermediate position is to place the bulk in Thelypteris (which can then be divided into subgenera and sections corresponding to the genera of other authors) but to separate out Phegopteris and Macrothelypteris.[2] Another choice is to divide the family into a half a dozen or so genera.[5]

Phylogeny of Thelypteriaceae[11][12]

The Pteridophyte Phylogeny Group classification of 2016 (PPG I) accepts 30 genera:[3]

Some species of Pronephrium were split off into other genera in 2021 as a result of a phylogenetic study of the family Thelypteridaceae:[15][16]

The extinct genus Holttumopteris from the Cenomanian aged Burmese amber has been suggested to have affinities with the family, but several important diagnostic characters are not preserved.[17]

References

[edit]
  1. ^ Thelypteridaceae Ching ex Pic. Serm. Tropicos.org. Missouri Botanical Garden. 24 Jan 2012
  2. ^ a b c "Thelypteridaceae". Flora of North America.
  3. ^ a b c d e f 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. S2CID 39980610.
  4. ^ Maarten J. M. Christenhusz; Xian-Chun Zhang & Harald Schneider (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.
  5. ^ a b c d Alan R. Smith; Kathleen M. Pryer; Eric Schuettpelz; Petra Korall; Harald Schneider & Paul G. Wolf (2006). "A classification for extant ferns" (PDF). Taxon. 55 (3): 705–731. doi:10.2307/25065646. JSTOR 25065646.
  6. ^ Christenhusz, Maarten J.M. & Chase, Mark W. (2014). "Trends and concepts in fern classification". Annals of Botany. 113 (9): 571–594. doi:10.1093/aob/mct299. PMC 3936591. PMID 24532607.
  7. ^ Collinson, M. E. (2001). "Cainozoic ferns and their distribution". Brittonia. 53 (2): 173–235. doi:10.1007/BF02812700. JSTOR 2666600. S2CID 19984401.
  8. ^ Samuli Lehtonen (2011). "Towards Resolving the Complete Fern Tree of Life" (PDF). PLOS ONE. 6 (10): e24851. Bibcode:2011PLoSO...624851L. doi:10.1371/journal.pone.0024851. PMC 3192703. PMID 22022365.
  9. ^ Carl J. Rothfels; Anders Larsson; Li-Yaung Kuo; Petra Korall; Wen- Liang Chiou; Kathleen M. Pryer (2012). "Overcoming Deep Roots, Fast Rates, and Short Internodes to Resolve the Ancient Rapid Radiation of Eupolypod II Ferns". Systematic Biology. 61 (1): 490–509. arXiv:cond-mat/0606434. doi:10.1093/sysbio/sys001. PMID 22223449.
  10. ^ a b Hassler, Michael & Schmitt, Bernd (January 2020). "Thelypteridaceae". Checklist of Ferns and Lycophytes of the World. 8.20. Retrieved 2020-01-11.
  11. ^ 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.
  12. ^ "Tree viewer: interactive visualization of FTOL". FTOL v1.3.0. 2022. Retrieved 12 December 2022.
  13. ^ Hassler, Michael & Schmitt, Bernd (January 2020). "Phegopteris". Checklist of Ferns and Lycophytes of the World. 8.20. Retrieved 2020-01-10.
  14. ^ "Phegopteris (C.Presl) Fée". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 2020-01-10.
  15. ^ Fawcett, S. & Smith, A.R. (2021), A Generic Classification of the Thelypteridaceae, Sida, Botanical Miscellany 59, Fort Worth: Botanical Research Institute of Texas Press
  16. ^ Fawcett, Susan; Smith, Alan R.; Sundue, Michael; Burleigh, J. Gordon; Sessa, Emily B.; Kuo, Li-Yaung; Chen, Cheng-Wei; Testo, Weston L.; Kessler, Michael & Barrington, David S. (2021), "A Global Phylogenomic Study of the Thelypteridaceae" (PDF), Systematic Botany, 46 (4): 891–915, doi:10.1600/036364421X16370109698650, S2CID 245512861, retrieved 2022-06-04
  17. ^ Regalado, Ledis; Schmidt, Alexander R.; Krings, Michael; Bechteler, Julia; Schneider, Harald; Heinrichs, Jochen (January 2018). "Fossil evidence of eupolypod ferns in the mid-Cretaceous of Myanmar". Plant Systematics and Evolution. 304 (1): 1–13. doi:10.1007/s00606-017-1439-2. ISSN 0378-2697. S2CID 21617872.
Retrieved from "https://en.wikipedia.org/w/index.php?title=Thelypteridaceae&oldid=1181866748"