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Leptosporangiate fern

November 24, 2023

The Polypodiidae, commonly called leptosporangiate ferns, formerly Leptosporangiatae, are one of four subclasses of ferns, the largest of these being the largest group of living ferns, including some 11,000 species worldwide.[2][3][4] The group has also been treated as the class Pteridopsida or Polypodiopsida,[5] although other classifications assign them a different rank.[6] Older names for the group include Filicidae and Filicales, although at least the "water ferns" (now the Salviniales) were then treated separately.

The leptosporangiate ferns are one of the four major groups of ferns, with the other three being the eusporangiate ferns comprising the marattioid ferns (Marattiidae, Marattiaceae), the horsetails (Equisetiidae, Equisetaceae), and whisk ferns and moonworts.[4][5] There are approximately 8465 species of living leptosporangiate ferns, compared with about 2070 for all other ferns, totalling 10535 species of ferns.[3] Almost a third of leptosporangiate fern species are epiphytes.[7]

These ferns are called leptosporangiate because their sporangia arise from a single epidermal cell and not from a group of cells as in eusporangiate ferns (a polyphyletic lineage). The mature sporangia have a wall that is just a single cell thick,[8] and are typically covered with a scale called the indusium, which can cover the whole sorus, forming a ring or cup around the sorus, or can also be strongly reduced to completely absent. Many leptosporangiate ferns have an annulus around the sporangium, which ejects the spores.[9]

Taxonomy edit

The leptosporangiate ferns were first recognized as a group, the "Leptosporangiateen", by Karl Ritter von Goebel in 1881, who placed the eusporangiate ferns with seed plants and vascular plants into a coeval "Eusporangiateen". As this classification artificially split the ferns, Christian Luerssen subdivided the homosporous ferns only into Eusporangiatae and Leptosporangiatae in 1884–9. The latter group was treated at a variety of ranks in subsequent systems of classification. The subclass "Polypodiidae" was first published and used for the homosporous leptosporangiate ferns by Cronquist, Takhtajan and Zimmermann in 1966, typified on Polypodium L.. Other contemporary classifications used the name "Filicidae" for this subclass.[10]

Smith et al. (2006) carried out the first higher-level classification of ferns based on molecular phylogenetics. They included heterosporous water ferns (Salviniales) (placed in a separate subclass by Cronquist et al. due to their highly modified morphology) within the leptosporangiate ferns, which they elevated to the rank of class as the Polypodiopsida (published by Cronquist et al. to include all ferns).[5]

The common ancestor of Salviniales, Cyatheales and Polypodiales went through a whole genome duplication.[11]

Later classifications renamed the group Polypodiidae, initially as a subclass of Equisetopsida sensu lato.[6] This subclass comprises leptosporangiate ferns as opposed to the remaining three subclasses which are informally referred to as eusporangiate ferns. The following diagram shows a likely phylogenic relationship between subclass Polypodiidae and the other Equisetopsida subclasses in that system[4]

In 2014, Christenhusz and Chase grouped all the fern subclasses together as Polypodiophyta[3] and in 2016 the Pteridophyte Phylogeny Group (PPG) adopted the class Polypodiopsida sensu lato for the four fern subclasses. The following cladogram shows the phylogenic relationship between the subclasses according to the PPG. The first three small subclasses being informally grouped as eusporangiate ferns, in contrast to the Polypodiidae or leptosporangiate ferns. Polypodiidae is shown as a sister group of Marattiidae.[1]

Subdivision edit

In both the Christenhusz and Chase, and the PPG classification, the extant Polypodiidae are divided into seven orders, 44 families, 300 genera, and an estimated 10,323 species.[1][3]

Christenhusz and Chase 2014 [3] Nitta et al. 2022[12] and Fern Tree of life[13]
Polypodiidae

Osmundales 1 family

Hymenophyllales 1 family

Gleicheniales 3 families

Schizaeales 3 families

Salviniales 2 families

Cyatheales 8 families

Polypodiales 6 suborders, 26 families

Phylogenetic relationships edit

The following phylogram shows a likely relationship between the other vascular plant classes and the leptosporangiate ferns. It was formerly unclear about the relationship between Equisetopsida, Psilotopsida, and Marattiopsida,[14][15][16] but recent studies have shown that Equisetopsida is most likely sister to Psilotopsida.

Tracheophyta

Lycopodiophytes (club mosses, spike mosses, quillworts)

Euphyllophytes

Spermatophytes (seed plants)

Ferns
Psilotopsida

Psilotales (whisk ferns)  

Ophioglossales (grapeferns etc.)

Equisetopsida

Equisetales (horsetails)  


Discussion of molecular classification edit

There has been some challenge to recent molecular studies, claiming that these provide a skewed view of the phylogenetic order because they do not take into account fossil representatives.[17] However, the molecular studies have clarified relations among families that had already been thought to be polyphyletic before the advent of molecular information but that were left in their polyphyletic ranks because there was not enough information to do otherwise.[18] The classification of ferns using these molecular studies, which have generally supported one another, reflects the best information available at present, because traditional morphological characters are not always informative in elucidating evolutionary relationships among ferns.[3]

Extinct families edit

The leptosporangiate ferns have a substantial fossil record. For example, fossils assigned to the Dicksoniaceae, a member of the Cyatheales, are known from the Lower Jurassic (201 to 175 million years ago).[19] A number of other extinct families have been described. They are not included in the classification systems used for extant ferns, and so most cannot be assigned to orders used in these systems. Taylor et al. (2009) use the order "Filicales", which corresponds to four Polypodiidae orders in more modern systems: Hymenophyllales, Gleicheniales, Schizaeales and Cyatheales. The unplaced families include:[20]

  • Anachoropteridaceae
  • Botryopteridaceae
  • Kaplanopteridaceae
  • Psalixochlaenaceae
  • Sermayaceae
  • Skaaripteridaceae
  • Tedeleaceae
  • Tempskyaceae

References edit

  1. ^ a b c Pteridophyte Phylogeny Group (2016).
  2. ^ Palmer, Jeffrey (2004), "The Plant Tree of Life: an Overview and Some Points of View", American Journal of Botany, 91 (10): 1437–45, doi:10.3732/ajb.91.10.1437, PMID 21652302
  3. ^ a b c d e f Christenhusz & Chase (2014).
  4. ^ a b c Christenhusz et al. (2011).
  5. ^ a b c Smith et al. (2006).
  6. ^ a b Chase & Reveal (2009).
  7. ^ Schuettpelz, Eric (2007). "Fern Phylogeny Inferred from 400 Leptosporangiate Species and Three Plastid Genes" (PDF). The Evolution and Diversification of Epiphytic Ferns (Doctoral dissertation). Duke University. Retrieved 2019-12-04.
  8. ^ Lack, Andrew J.; Evans, David E. (2005). Plant Biology. ISBN 9780415356435.
  9. ^ Llorens, C.; Argentina, M.; Rojas, N.; Westbrook, J.; Dumais, J.; Noblin, X. (2016). "The fern cavitation catapult: Mechanism and design principles". Journal of the Royal Society Interface. 13 (114). doi:10.1098/rsif.2015.0930. PMC 4759797. PMID 26763327.
  10. ^ Cronquist, Arthur; Takhtajan, Armen & Zimmermann, Walter (April 1966). "On the Higher Taxa of Embryobionta". Taxon. 15 (4): 129–134. doi:10.2307/1217531. JSTOR 1217531.
  11. ^ Li, Fay-Wei; Brouwer, Paul; Carretero-Paulet, L.; et al. (2018). "Fern genomes elucidate land plant evolution and cyanobacterial symbioses". Nature Plants. 4 (7): 460–472. doi:10.1038/s41477-018-0188-8. PMC 6786969. PMID 29967517.
  12. ^ 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. doi:10.3389/fpls.2022.909768. PMC 9449725. PMID 36092417.
  13. ^ "Tree viewer: interactive visualization of FTOL". FTOL v1.3.0. 2022. Retrieved 12 December 2022.
  14. ^ 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.
  15. ^ Hardeep S. Rai; Sean W. Graham (2010). "Utility of a large, multigene plastid data set in inferring higher-order relationships in ferns and relatives (Monilophytes)" (PDF). American Journal of Botany. 97 (9): 1444–1456. doi:10.3732/ajb.0900305. PMID 21616899.
  16. ^ Kathleen M. Pryer; Eric Schuettpelz (2009). "Ferns" (PDF). In S. Blair Hedges; Sudhir Kumar (eds.). The Timetree of Life. Oxford Biology.
  17. ^ Rothwell, G. W. & Nixon, K. C. (2006). "How does the inclusion of fossil data change our conclusions about the phylogenetic history of euphyllophytes". International Journal of Plant Sciences. 167 (3): 737–749. doi:10.1086/503298. S2CID 86172890.
  18. ^ Kramer, K. U. (1990). Notes on the Higher Level Classification of the Recent Ferns. The Families and Genera of Vascular Plants: Pteridophytes and Gymnosperms. K. Kubitzki, K. U. Kramer and P. S. Green. New York, Springer-Verlag. 1: 49-52
  19. ^ Taylor, Taylor & Krings (2009), pp. 464.
  20. ^ Taylor, Taylor & Krings (2009), pp. 436–476.

Bibliography edit

  • Chase, Mark W.; Reveal, James L. (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.
  • Christenhusz, M.J.M.; Zhang, X.C.; Schneider, H. (18 February 2011). "A linear sequence of extant families and genera of lycophytes and ferns". Phytotaxa. 19 (1): 7. doi:10.11646/phytotaxa.19.1.2.
  • Christenhusz, Maarten J.M.; Chase, Mark W. (2014). "Trends and concepts in fern classification". Annals of Botany. 113 (4): 571–594. doi:10.1093/aob/mct299. PMC 3936591. PMID 24532607.
  • 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.
  • 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.
  • Taylor, T.N.; Taylor, E.L. & Krings, M. (2009). Paleobotany, The Biology and Evolution of Fossil Plants (2nd ed.). Amsterdam; Boston: Academic Press. ISBN 978-0-12-373972-8.</ref>

November 24, 2023
leptosporangiate, fern, polypodiidae, commonly, called, leptosporangiate, ferns, formerly, leptosporangiatae, four, subclasses, ferns, largest, these, being, largest, group, living, ferns, including, some, species, worldwide, group, also, been, treated, class,. The Polypodiidae commonly called leptosporangiate ferns formerly Leptosporangiatae are one of four subclasses of ferns the largest of these being the largest group of living ferns including some 11 000 species worldwide 2 3 4 The group has also been treated as the class Pteridopsida or Polypodiopsida 5 although other classifications assign them a different rank 6 Older names for the group include Filicidae and Filicales although at least the water ferns now the Salviniales were then treated separately Leptosporangiate fernPteridium aquilinumScientific classificationKingdom PlantaeClade TracheophytesDivision PolypodiophytaClass PolypodiopsidaSubclass PolypodiidaeCronquist Takht amp W Zimm 1 OrdersOsmundales Hymenophyllales Gleicheniales Schizaeales Salviniales Cyatheales PolypodialesThe leptosporangiate ferns are one of the four major groups of ferns with the other three being the eusporangiate ferns comprising the marattioid ferns Marattiidae Marattiaceae the horsetails Equisetiidae Equisetaceae and whisk ferns and moonworts 4 5 There are approximately 8465 species of living leptosporangiate ferns compared with about 2070 for all other ferns totalling 10535 species of ferns 3 Almost a third of leptosporangiate fern species are epiphytes 7 These ferns are called leptosporangiate because their sporangia arise from a single epidermal cell and not from a group of cells as in eusporangiate ferns a polyphyletic lineage The mature sporangia have a wall that is just a single cell thick 8 and are typically covered with a scale called the indusium which can cover the whole sorus forming a ring or cup around the sorus or can also be strongly reduced to completely absent Many leptosporangiate ferns have an annulus around the sporangium which ejects the spores 9 Contents 1 Taxonomy 1 1 Subdivision 1 2 Phylogenetic relationships 2 Discussion of molecular classification 3 Extinct families 4 References 5 BibliographyTaxonomy editThe leptosporangiate ferns were first recognized as a group the Leptosporangiateen by Karl Ritter von Goebel in 1881 who placed the eusporangiate ferns with seed plants and vascular plants into a coeval Eusporangiateen As this classification artificially split the ferns Christian Luerssen subdivided the homosporous ferns only into Eusporangiatae and Leptosporangiatae in 1884 9 The latter group was treated at a variety of ranks in subsequent systems of classification The subclass Polypodiidae was first published and used for the homosporous leptosporangiate ferns by Cronquist Takhtajan and Zimmermann in 1966 typified on Polypodium L Other contemporary classifications used the name Filicidae for this subclass 10 Smith et al 2006 carried out the first higher level classification of ferns based on molecular phylogenetics They included heterosporous water ferns Salviniales placed in a separate subclass by Cronquist et al due to their highly modified morphology within the leptosporangiate ferns which they elevated to the rank of class as the Polypodiopsida published by Cronquist et al to include all ferns 5 The common ancestor of Salviniales Cyatheales and Polypodiales went through a whole genome duplication 11 Later classifications renamed the group Polypodiidae initially as a subclass of Equisetopsida sensu lato 6 This subclass comprises leptosporangiate ferns as opposed to the remaining three subclasses which are informally referred to as eusporangiate ferns The following diagram shows a likely phylogenic relationship between subclass Polypodiidae and the other Equisetopsida subclasses in that system 4 Equisetopsida MarchantiidaeBryidaeAnthocerotidaeLycopodiidaeEquisetidaeOphioglossidaeMarattiidaePolypodiidaeCycadidaeGinkgoidaeGnetidaePinidaeMagnoliidae bryophytes lycopodiophytes monilophytes gymnosperms angiospermsIn 2014 Christenhusz and Chase grouped all the fern subclasses together as Polypodiophyta 3 and in 2016 the Pteridophyte Phylogeny Group PPG adopted the class Polypodiopsida sensu lato for the four fern subclasses The following cladogram shows the phylogenic relationship between the subclasses according to the PPG The first three small subclasses being informally grouped as eusporangiate ferns in contrast to the Polypodiidae or leptosporangiate ferns Polypodiidae is shown as a sister group of Marattiidae 1 Polypodiopsida EquisetidaeOphioglossidaeMarattiidaePolypodiidaeSubdivision edit In both the Christenhusz and Chase and the PPG classification the extant Polypodiidae are divided into seven orders 44 families 300 genera and an estimated 10 323 species 1 3 Christenhusz and Chase 2014 3 Nitta et al 2022 12 and Fern Tree of life 13 Polypodiidae Osmundales 1 familyHymenophyllales 1 familyGleicheniales 3 familiesSchizaeales 3 familiesSalviniales 2 familiesCyatheales 8 familiesPolypodiales 6 suborders 26 families OsmundalesHymenophyllalesGleichenialesSchizaealesSalvinialesCyathealesPolypodialesPhylogenetic relationships edit The following phylogram shows a likely relationship between the other vascular plant classes and the leptosporangiate ferns It was formerly unclear about the relationship between Equisetopsida Psilotopsida and Marattiopsida 14 15 16 but recent studies have shown that Equisetopsida is most likely sister to Psilotopsida Tracheophyta Lycopodiophytes club mosses spike mosses quillworts Euphyllophytes Spermatophytes seed plants Ferns Psilotopsida Psilotales whisk ferns nbsp Ophioglossales grapeferns etc Equisetopsida Equisetales horsetails nbsp Marattiopsida Marattiales nbsp Polypodiopsida Osmundales nbsp Hymenophyllales filmy ferns nbsp Gleicheniales nbsp SchizaealesSalviniales heterosporous Cyatheales tree ferns nbsp Polypodiales nbsp EusporangiateFerns LeptosporangiateFernsDiscussion of molecular classification editThere has been some challenge to recent molecular studies claiming that these provide a skewed view of the phylogenetic order because they do not take into account fossil representatives 17 However the molecular studies have clarified relations among families that had already been thought to be polyphyletic before the advent of molecular information but that were left in their polyphyletic ranks because there was not enough information to do otherwise 18 The classification of ferns using these molecular studies which have generally supported one another reflects the best information available at present because traditional morphological characters are not always informative in elucidating evolutionary relationships among ferns 3 Extinct families editThe leptosporangiate ferns have a substantial fossil record For example fossils assigned to the Dicksoniaceae a member of the Cyatheales are known from the Lower Jurassic 201 to 175 million years ago 19 A number of other extinct families have been described They are not included in the classification systems used for extant ferns and so most cannot be assigned to orders used in these systems Taylor et al 2009 use the order Filicales which corresponds to four Polypodiidae orders in more modern systems Hymenophyllales Gleicheniales Schizaeales and Cyatheales The unplaced families include 20 Anachoropteridaceae Botryopteridaceae Kaplanopteridaceae Psalixochlaenaceae Sermayaceae Skaaripteridaceae Tedeleaceae TempskyaceaeReferences edit a b c Pteridophyte Phylogeny Group 2016 Palmer Jeffrey 2004 The Plant Tree of Life an Overview and Some Points of View American Journal of Botany 91 10 1437 45 doi 10 3732 ajb 91 10 1437 PMID 21652302 a b c d e f Christenhusz amp Chase 2014 a b c Christenhusz et al 2011 a b c Smith et al 2006 a b Chase amp Reveal 2009 Schuettpelz Eric 2007 Fern Phylogeny Inferred from 400 Leptosporangiate Species and Three Plastid Genes PDF The Evolution and Diversification of Epiphytic Ferns Doctoral dissertation Duke University Retrieved 2019 12 04 Lack Andrew J Evans David E 2005 Plant Biology ISBN 9780415356435 Llorens C Argentina M Rojas N Westbrook J Dumais J Noblin X 2016 The fern cavitation catapult Mechanism and design principles Journal of the Royal Society Interface 13 114 doi 10 1098 rsif 2015 0930 PMC 4759797 PMID 26763327 Cronquist Arthur Takhtajan Armen amp Zimmermann Walter April 1966 On the Higher Taxa of Embryobionta Taxon 15 4 129 134 doi 10 2307 1217531 JSTOR 1217531 Li Fay Wei Brouwer Paul Carretero Paulet L et al 2018 Fern genomes elucidate land plant evolution and cyanobacterial symbioses Nature Plants 4 7 460 472 doi 10 1038 s41477 018 0188 8 PMC 6786969 PMID 29967517 Nitta Joel H Schuettpelz Eric Ramirez Barahona Santiago Iwasaki Wataru et al 2022 An Open and Continuously Updated Fern Tree of Life Frontiers in Plant Science 13 doi 10 3389 fpls 2022 909768 PMC 9449725 PMID 36092417 Tree viewer interactive visualization of FTOL FTOL v1 3 0 2022 Retrieved 12 December 2022 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 Hardeep S Rai Sean W Graham 2010 Utility of a large multigene plastid data set in inferring higher order relationships in ferns and relatives Monilophytes PDF American Journal of Botany 97 9 1444 1456 doi 10 3732 ajb 0900305 PMID 21616899 Kathleen M Pryer Eric Schuettpelz 2009 Ferns PDF In S Blair Hedges Sudhir Kumar eds The Timetree of Life Oxford Biology Rothwell G W amp Nixon K C 2006 How does the inclusion of fossil data change our conclusions about the phylogenetic history of euphyllophytes International Journal of Plant Sciences 167 3 737 749 doi 10 1086 503298 S2CID 86172890 Kramer K U 1990 Notes on the Higher Level Classification of the Recent Ferns The Families and Genera of Vascular Plants Pteridophytes and Gymnosperms K Kubitzki K U Kramer and P S Green New York Springer Verlag 1 49 52 Taylor Taylor amp Krings 2009 pp 464 Taylor Taylor amp Krings 2009 pp 436 476 Bibliography editChase Mark W Reveal James L 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 Christenhusz M J M Zhang X C Schneider H 18 February 2011 A linear sequence of extant families and genera of lycophytes and ferns Phytotaxa 19 1 7 doi 10 11646 phytotaxa 19 1 2 Christenhusz Maarten J M Chase Mark W 2014 Trends and concepts in fern classification Annals of Botany 113 4 571 594 doi 10 1093 aob mct299 PMC 3936591 PMID 24532607 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 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 Taylor T N Taylor E L amp Krings M 2009 Paleobotany The Biology and Evolution of Fossil Plants 2nd ed Amsterdam Boston Academic Press ISBN 978 0 12 373972 8 lt ref gt Retrieved from https en wikipedia org w index php title Leptosporangiate fern amp oldid 1180500436, wikipedia, wiki, book, books, library,

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