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Leptosporangiate fern
Pteridium aquilinum 3 BOGA.jpg
Pteridium aquilinum
Scientific classification e
Kingdom: Plantae
Clade: Tracheophytes
Class: Polypodiopsida
Subclass: Polypodiidae
Cronquist, Takht. & W.Zimm.
Orders
  • Osmundales
  • Hymenophyllales
  • Gleicheniales
  • Schizaeales
  • Salviniales
  • Cyatheales
  • Polypodiales

The Polypodiidae, commonly called leptosporangiate ferns, formerly Leptosporangiatae, are one of four subclasses of ferns, and the largest of these, being the largest group of living ferns, including some 11,000 species worldwide. The group has also been treated as the class Pteridopsida or Polypodiopsida, although other classifications assign them a different rank. 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. There are approximately 8465 species of living leptosporangiate ferns, compared with about 2070 for all other ferns, totalling 10535 species of ferns. Almost a third of leptosporangiate fern species are epiphytes.

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 sporangia 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.

Taxonomy

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.

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).

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

Later classifications renamed the group Polypodiidae, initially as a subclass of Equisetopsida sensu lato. 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

Equisetopsida








Marchantiidae  

















Bryidae  















Anthocerotidae  













Lycopodiidae  











Equisetidae  





Ophioglossidae  





Marattiidae  





Polypodiidae  









Cycadidae  



Ginkgoidae  





Gnetidae  



Pinidae  







Magnoliidae  










lycopodiophytes
monilophytes

In 2014, Christenhusz and Chase grouped all the fern subclasses together as Polypodiophyta 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.

Polypodiopsida

Equisetidae




Ophioglossidae




Marattiidae



Polypodiidae






Subdivision

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.

These seven orders are phylogenetically related as follows (Christenhusz and Chase 2014 in italics, where it differs):

Polypodiidae

Osmundales 1 family




Hymenophyllales 1 family




Gleicheniales 3 families




Schizaeales 3 families (1 family, 3 subfamilies)




Salviniales 2 families




Cyatheales 8 families (1 family, 8 subfamilies)



Polypodiales 6 suborders, 26 families (0 suborders, 8 families)









Phylogenetic relationships

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, 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)  





Marattiopsida

Marattiales  



Polypodiopsida

Osmundales  




Hymenophyllales (filmy ferns)  




Gleicheniales  




Schizaeales  




Salviniales (heterosporous)  




Cyatheales (tree ferns)  



Polypodiales  












Eusporangiate
Ferns
Leptosporangiate
Ferns


Extinct families

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 174 million years ago). 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:

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

See also

Kids robot.svg In Spanish: Helechos leptosporangiados para niños

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