Marsh fern family
Temporal range:
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

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

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)

Cystopteridaceae

Rhachidosoraceae

Diplaziopsidaceae

Aspleniaceae

Hemidictyaceae

Thelypteridaceae

Woodsiaceae

Onocleaceae

Blechnaceae

Athyriaceae

Genera

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; 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]

Phegopteridoideae

Macrothelypteris

Phegopteris

Pseudophegopteris

Thelypteridoideae
Thelypterideae

Thelypteris

Amauropelteae

Coryphopteris

Metathelypteris

Amauropelta

Oreopteris

Steiropteris

Leptogrammeae

Cyclogramma

Stegnogramma

Leptogramma

Meniscieae

Steiropteris species-group 2

Goniopteridinae

Goniopteris

Menisciinae

Cyclosorus

Ampelopteris

Mesophlebion

Meniscium

Pseudocyclosorinae

Pelazoneuron

Pakau

Pelazoneuron species-group 2

Glaphyropteridopsis

Chingia

Plesioneuron

Menisciopsis

Grypothrix

Mesopteris

Plesioneuron archboldiae

Strophocaulon

Pneumatopteris

Reholttumia

Pronephrium

Sphaerostephanos

Trigonospora

Pseudocyclosorus

Amblovenatum

Sphaerostephanos elatus

Abacopteris

Christella

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

  1. ↑ Thelypteridaceae Ching ex Pic. Serm. Tropicos.org. Missouri Botanical Garden. 24 Jan 2012
  2. 1 2 3 "Thelypteridaceae". Flora of North America.
  3. 1 2 3 4 5 6 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. 1 2 3 4 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. 1 2 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.
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