Polypodiales

Polypodiales
Temporal range: Early Cretaceous–Recent
California Arena Point fern.jpg
Polypodium californicum
Scientific classification e
Kingdom: Plantae
Clade: Tracheophytes
Division: Polypodiophyta
Class: Polypodiopsida
Subclass: Polypodiidae
Order: Polypodiales
Link (1833)
Suborders[1]

6. See text

The order Polypodiales encompasses the major lineages of polypod ferns, which comprise more than 80% of today's fern species. They are found in many parts of the world including tropical, semitropical and temperate areas.

Description

Polypodiales are unique in bearing sporangia with a vertical annulus interrupted by the stalk and stomium.[2] These sporangial characters were used by Johann Jakob Bernhardi to define a group of ferns he called the "Cathetogyratae";[3] the Pteridophyte Phylogeny Group has suggested reviving this name as the informal term cathetogyrates, to replace the ambiguously circumscribed term "polypods" when referring to the Polypodiales.[1] The sporangia are born on stalks 1–3 cells thick and are often long-stalked.[2] (In contrast, the Hymenophyllales have a stalk composed of four rows of cells.)[4] The sporangia do not reach maturity simultaneously. Many groups in the order lack indusia, but when present, they are attached either along the edge of the indusium or in its center.[2]

Both Polypodiales and Cyatheales differ from other ferns in having a photoreceptor called a neochrome, which allows them to perform photosynthesis better in low-light conditions, such as in the shadows on the forest floor. The common ancestor of the two groups appears to have derived the neochrome via horizontal gene transfer from a hornwort.[5]

Their gametophytes are green, usually heart-shaped, and grow at the surface[2] (rather than underground, as in Ophioglossales).[6]

Taxonomy

The order Polypodiales was first described by Link in 1833.[1] The circumscription of the order has changed over time as ferns have been classified in many different ways (see the review by Christenhusz and Chase, 2014).[7] Smith et al. (2006) carried out the first higher-level pteridophyte classification published in the molecular phylogenetic era.[8] They referred to the ferns (now including horsetails) as monilophytes, dividing them into four groups, with the vast majority of species being placed in a taxon they called "Polypodiopsida". The four-fold grouping has persisted through subsequent systems, despite changes in nomenclature.[9][10][7][1] Polypodiopsida is now used for all ferns (sensu lato),[1] with Smith et al.'s group being subclass Polypodiidae. This group, which includes Polypodiales, is also informally known as the leptosporangiate ferns, while the remaining three groups (subclasses) are referred to as eusporangiate ferns. The Polypodiidae have been divided into seven orders, Polypodiales being the largest. The phylogenetic position of Polypodiales in relation to the other orders of Polypodiidae is shown in the following cladogram.[1]

Subclass Polypodiidae

Osmundales

Hymenophyllales

Gleicheniales

Schizaeales

Salviniales

Cyatheales

Polypodiales (6 suborders, 26 families)

Subdivision

The division of the Polypodiales into families has changed somewhat between the pioneering work of Smith et al. (2006) and the Pteridophyte Phylogeny Group's classification of 2016, with a general increase in the number of divisions recognized, albeit sometimes at different ranks. The table below summarizes four systems; families are listed alphabetically within three broad groups. Although the same families are used in more than one system, circumscriptions may differ. Christenhusz and Chase in 2014 used a very broad circumscription of Aspleniaceae and Polypodiaceae, reducing families used in other systems to subfamilies.

Comparison of alternative subdivisions of Polypodiales
Smith et al. (2006)[8] Christenhusz et al. (2011)[10] Christenhusz & Chase (2014)[7] PPG I (2016)[1]
Basal families Cystodiaceae Cystodiaceae Cystodiaceae
Dennstaedtiaceae Dennstaedtiaceae Dennstaedtiaceae Dennstaedtiaceae
Lindsaeaceae Lindsaeaceae Lindsaeaceae Lindsaeaceae
Lonchitidaceae Lonchitidaceae Lonchitidaceae
Pteridaceae Pteridaceae Pteridaceae Pteridaceae
Saccolomataceae Saccolomataceae Saccolomataceae Saccolomataceae
Aspleniineae
eupolypods II
(Aspleniaceae)
Aspleniaceae Aspleniaceae Aspleniaceae: Asplenioideae Aspleniaceae
Athyriaceae Aspleniaceae: Athyrioideae Athyriaceae
Blechnaceae Blechnaceae Aspleniaceae: Blechnoideae Blechnaceae
Cystopteridaceae Aspleniaceae: Cystopteridoideae Cystopteridaceae
Desmophlebiaceae
Diplaziopsidaceae Aspleniaceae: Diplaziopsidoideae Diplaziopsidaceae
Hemidictyaceae
Onocleaceae Onocleaceae Onocleaceae
Rhachidosoraceae Aspleniaceae: Rhachidosoroideae Rhachidosoraceae
Thelypteridaceae Thelypteridaceae Aspleniaceae: Thelypteridoideae Thelypteridaceae
Woodsiaceae Woodsiaceae Aspleniaceae: Woodsioideae Woodsiaceae
Polypodiineae
eupolypods I
(Polypodiaceae)
Davalliaceae Davalliaceae Polypodiaceae: Davallioideae Davalliaceae
Polypodiaceae: Didymochlaenoideae Didymochlaenaceae
Dryopteridaceae Dryopteridaceae Polypodiaceae: Dryopteridoideae Dryopteridaceae
Hypodematiaceae Polypodiaceae: Hypodematioideae Hypodematiaceae
Lomariopsidaceae Lomariopsidaceae Polypodiaceae: Lomariopsidoideae Lomariopsidaceae
Nephrolepidaceae Nephrolepidaceae
Oleandraceae Oleandraceae Polypodiaceae: Oleandroideae Oleandraceae
Polypodiaceae Polypodiaceae Polypodiaceae: Polypodioideae Polypodiaceae
Tectariaceae Tectariaceae Polypodiaceae: Tectarioideae Tectariaceae

Smith et al. (2006) divided the Polypodiales into fifteen families,[8] a practice continued in their 2008 revision,[11] with members of the eupolypods placed in two unranked clades. The families are listed in the table. While many of these families had previously been recognized with similar circumscriptions, the authors noted that Dryopteridaceae was more narrowly bounded than in historical circumscriptions, which had included their Tectariaceae, Onocleaceae and Woodsiaceae. The circumscription of Lomariopsidaceae changed dramatically, with most historical genera of that family (except Lomariopsis and Thysanosoria) being moved to Dryopteridaceae, while Cyclopeltis and Nephrolepis were added. Saccolomataceae were removed from the dennstaedtioids. Cystodium was tentatively placed in Lindsaeaceae, away from its historical position with the tree ferns. Woodsiaceae was acknowledged to be of uncertain circumscription and perhaps paraphyletic; the inclusion of Hypodematium, Didymochlaena, and Leucostegia perhaps also rendering Dryopteridaceae paraphyletic. The grammitids were included in Polypodiaceae to render that family monophyletic.[8]

The linear sequence of Christenhusz et al. (2011), intended for compatibility with the classification of Chase and Reveal (2009),[9] incorporated new phylogenetic evidence to make several changes at the familial level, resulting in an expansion to 23 families. Lonchitis and Cystodium were removed from the Lindsaeaceae and incorporated into new families, Lonchitidaceae and Cystodiaceae respectively. Within eupolypods I, Woodsiaceae proved to be paraphyletic and was reduced to the genera Cheilanthopsis, Hymenocystis, and Woodsia, while the remainder of its genera were removed to Cystopteridaceae, Diplaziopsidaceae, Rhachidosoraceae, Athyriaceae, and Hemidictyaceae. Within eupolypods II, Nephrolepis was placed in a new family, the Nephrolepidaceae, due to uncertainty in its phylogenetic placement, while Hypodematiaceae was split from Dryopteridaceae to contain the three problematic genera mentioned by Smith et al.[10]

The classification of Christenhusz and Chase (2014) dramatically reduced the number of families recognized in this order to eight by "lumping", reducing many families to subfamilies and expanding the circumscription of Polypodiaceae and Aspleniaceae to encompass all of eupolypods I and eupolypods II, respectively. Former families became subfamilies (see the table above). The former Hemidictyaceae were included in the Asplenioideae, and the Onocleaceae in the Blechnoideae. In the new Polypodiaceae, Didymochlaena was placed in its own subfamily, Didymochlaenoideae.[7]

The PPG I classification (2016) used a process intermediate between the two previous approaches, by introducing a new rank, that of suborder, and organising 26 families (in some cases very narrowly circumscribed) into six suborders, largely returning to the families set out by Christenhusz et al. in 2011. In lieu of the expansion of Aspleniaceae and Polypodiaceae, eupolypods I and II were recognized and named as suborders:[1][7]

  • Saccolomatineae includes the single family Saccolomataceae.
  • Lindsaeinae corresponds to the Lindseaceae of Smith et al., and includes the Cystodiaceae, Lindsaeaceae, and Lonchitidaceae. It is probably not monophyletic.
  • Pteridineae includes the single family Pteridaceae.
  • Dennstaedtiineae includes the single family Dennstaedtiaceae.
  • Aspleniinae (formerly eupolypods I) includes the families Cystopteridaceae, Rhachidosoraceae, Diplaziopsidaceae, Desmophlebiaceae (containing only Desmophlebium), Hemidictyaceae, Aspleniaceae, Woodsiaceae, Onocleaceae, Blechnaceae, Athyriaceae, and Thelypteridaceae.
  • Polypodiineae (formerly eupolypods II) includes the families Didymochlaenaceae (containing only Didymochlaena), Hypodematiaceae, Dryopteridaceae, Lomariopsidaceae, Nephrolepidaceae, Tectariaceae, Oleandraceae, Davalliaceae, and Polypodiaceae.

Phylogeny

Pteridophyte Phylogeny Group 2016[1] Nitta et al. 2022[12] and Fern Tree of life[13]
Polypodiales

Saccolomataceae

Cystodiaceae

Lonchitidaceae

Lindsaeaceae

Pteridaceae

Dennstaedtiaceae

eupolypods
Aspleniineae

Cystopteridaceae

Rhachidosoraceae

Diplaziopsidaceae

Desmophlebiaceae

Hemidictyaceae

Aspleniaceae

Thelypteridaceae

Woodsiaceae

Athyriaceae

Onocleaceae

Blechnaceae

eupolypods II
Polypodiineae

Didymochlaenaceae

Hypodematiaceae

Dryopteridaceae

Nephrolepidaceae

Lomariopsidaceae

Tectariaceae

Oleandraceae

Davalliaceae

Polypodiaceae

eupolypods I

Saccolomataceae

Cystodiaceae

Lonchitidaceae

Lindsaeaceae

Pteridaceae

Dennstaedtiaceae

eupolypods
Aspleniineae

Cystopteridaceae

Rhachidosoraceae

Diplaziopsidaceae

Desmophlebiaceae

Hemidictyaceae

Aspleniaceae

Thelypteridaceae

Woodsiaceae

Onocleaceae

Blechnaceae

Athyriaceae

eupolypods II
Polypodiineae

Hypodematiaceae

Didymochlaenaceae

Dryopteridaceae

Lomariopsidaceae

Nephrolepidaceae

Tectariaceae

Oleandraceae

Davalliaceae

Polypodiaceae

eupolypods I

Obsolete families

Now-obsolete families of Polypodiales include:

  • Drynariaceae - now in Polypodiaceae
  • Grammitidaceae - now in Polypodiaceae
  • Gymnogrammitidaceae - now in Polypodiaceae
  • Loxogrammaceae - now in Polypodiaceae
  • Platyceriaceae - now in Polypodiaceae
  • Pleursoriopsidaceae - now in Polypodiaceae
  • Vittariaceae - now in Pteridaceae

Evolution

Polypodiales may be regarded as one of the most evolutionarily advanced orders of monilophytes (ferns), based on recent genetic analysis. They arose and diversified about 100 million years ago, probably subsequent to the diversification of the angiosperms.[14]

References

  1. ^ a b c d e f g h i Pteridophyte Phylogeny Group 2016.
  2. ^ a b c d Smith et al. 2006, p. 713.
  3. ^ Bernhardi 1806, pp. 9–12.
  4. ^ Smith et al. 2006, p. 711.
  5. ^ Li et al. 2014.
  6. ^ Smith et al. 2006, p. 710.
  7. ^ a b c d e Christenhusz & Chase 2014.
  8. ^ a b c d Smith et al. 2006.
  9. ^ a b Chase & Reveal 2009.
  10. ^ a b c Christenhusz et al. 2011.
  11. ^ Smith et al. 2008.
  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: 909768. 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. ^ Schneider et al. 2004.

Bibliography

  • Bernhardi, J. Jacob (1806). "Dritter Versuch einer Anordnung der Farrnkräuter". Neues Journal für die Botanik (in German). 1 (2): 1–50.
  • 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.
  • Christenhusz, Maarten JM & Byng, J. W. (2016). "The number of known plants species in the world and its annual increase". Phytotaxa. 261 (3): 201–217. doi:10.11646/phytotaxa.261.3.1.
  • Lehtonen, Samuli (2011). "Towards Resolving the Complete Fern Tree of Life". PLoS ONE. 6 (10): e24851. Bibcode:2011PLoSO...624851L. doi:10.1371/journal.pone.0024851. PMC 3192703. PMID 22022365.
  • Pryer, Kathleen M.; Schneider, Harald; Smith, Alan R.; Cranfill, Raymond; Wolf, Paul G.; Hunt, Jeffrey S.; Sipes, Sedonia D. (2001). "Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants". Nature. 409 (6820): 618–622. Bibcode:2001Natur.409..618S. doi:10.1038/35054555. PMID 11214320. S2CID 4367248.
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  • Ranker, Tom A.; Haufler, Christopher H., eds. (2008). Biology and Evolution of Ferns and Lycophytes. Cambridge University Press. ISBN 978-0-521-87411-3.
  • Schneider, Harald; Schuettpelz, Eric; Pryer, Kathleen M.; Cranfill, Raymond; Magallón, Susana; Lupia, Richard (1 April 2004). "Ferns diversified in the shadow of angiosperms". Nature. 428 (6982): 553–557. Bibcode:2004Natur.428..553S. doi:10.1038/nature02361. PMID 15058303.
  • Schneider, Harald; Smith, Alan R.; Pryer, Kathleen M. (1 July 2009). "Is Morphology Really at Odds with Molecules in Estimating Fern Phylogeny?". Systematic Botany. 34 (3): 455–475. doi:10.1600/036364409789271209. S2CID 85855934.
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