Polystichum

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Polystichum
Polystichum setiferum0.jpg
Polystichum setiferum
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Division: Polypodiophyta
Class: Polypodiopsida
Order: Polypodiales
Suborder: Polypodiineae
Family: Dryopteridaceae
Subfamily: Dryopteridoideae
Genus: Polystichum
Roth [1]
Species

See text.

Synonyms [2]
  • AcropeltaT.Nakai
  • AdenoderrisJ.Sm.
  • AetopteronEhrh. ex House
  • HemesteumH.Lév.
  • HypopeltisMichx.
  • PapuapterisC. Chr.
  • PhanerophlebiaC. Presl
  • PlecosorusFée
  • SorolepidiumChrist

Polystichum is a genus of ferns in the family Dryopteridaceae, subfamily Dryopteridoideae, according to the Pteridophyte Phylogeny Group classification of 2016 (PPG I). [1] The genus has about 500 species and has a cosmopolitan distribution. [1] [3] The highest diversity is in eastern Asia, with about 208 species in China alone; [3] the region from Mexico to Brazil has at least 100 additional species; Africa (at least 17 species), North America (at least 18 species), and Europe (at least 5 species) have much lower diversity. Polystichum species are terrestrial or rock-dwelling ferns of warm-temperate and montane-tropical regions (a few species grow in alpine regions). They are often found in disturbed habitats such as road cuts, talus slopes, and stream banks. [4]

Contents

Description

Many ferns of this genus have stout, slowly creeping rootstocks that form a crown, with a vase-like ring of evergreen fronds 30 to 200 centimetres (10 to 80 in) long. The sori are round, with a circular indusium, except in South American species which lack an indusium. [5] The stipes have prominent scales and often have hair-like cilia, but they lack any true hairs. [5] The genus differs from the well-known and allied fern genus Dryopteris in the indusium being circular, not reniform, and in having the leaf segments with auricles—asymmetrical blades where one side of the segment is much longer than the other at the base.

Apomixis

Apomixis, the development of an embryo without the occurrence of fertilization, is particularly common among ferns. Apomixis evolved several times independently in three different clades of polystichoid ferns. [6]

Taxonomy

Polystichum is one of the 10 largest fern genera and is grouped within the Dryopteridaceae. [7] Polystichum s.l. is well defined as its own monophyletic group, including species from the genera Cyrtomidictyum, Cyrtogonellum, Cyrtomium, and Phanerophlebia. [8] Research concerning taxonomy within Polystichum s.s. is ongoing, with high levels of hybridization, allopolyploidy, and apomixis making distinctions difficult. [9] Based on genetic analysis Little & Barrington (2003) originally defined a monophyletic Polystichum s.s. by removing Cyrtomium as its own genus. [4] It was further separated by Li et al. (2008) into a separate clade along with Phanerophlebia. [10]

Selected species

The genus has a large number of species. The PPG I classification suggested that there were about 500 species; [1] as of February 2020, the Checklist of Ferns and Lycophytes of the World listed 397 species and 58 hybrids, noting that "many undescribed species remain". [11] The genus Polystichum includes, but is not limited to, the following species. In this list, a species name preceded by (=) is considered to be a synonym of the accepted species name above it.

Polystichum munitum Polystichum munitum 1.jpg
Polystichum munitum

Hybrids

Hybridisation is frequent in the genus, with several named hybrids, including:-

Former species

Species that were at one time considered part of the genus Polystichum, but are now categorized elsewhere, include:

Distribution

With species in six continents and many islands, Polystichum is remarkable for its global spread. Polystichum ferns first emerged in Asia during the late Eocene, around 49 million years ago. [3] During this period there were high temperatures across the globe, which may have contributed to the diversification of flora. [45]

Polystichum's spread to the New World took place during the late Eocene to early Oligocene. The most likely dispersal method was across a paratropical forest on a Pacific Ocean land bridge, such as the Bering Land Bridge. [3] For a period before the height of the Ice Age temperatures froze enough seawater to lower the sea level but still allowed a forest to grow in Northern latitudes. From there Polystichum was able to spread through North American and into Central America.

Original theories described Polystichum spreading further into South America from Central America, but recent research has shown that South American Polystichum instead spread through long-distance dispersal from Australia. Genetic study has revealed close evolutionary relationships between Polystichum species in these two areas. Both Austral and Austral South American species lack and indusium. Austral Polystichum lack cilia, while South American species have marginal cilia. [46]

Hawaiian Polystichum also spread through long-distance dispersal, with two separate dispersal events leading to the three Polystichum now observed in Hawaii. [9]

Ecology

Polystichum species are used as food plants by the larvae of some Lepidoptera species including Pharmacis fusconebulosa . Specimens of some of these can be found at the Royal Botanic Gardens, Sydney.

Cultivation

Several species are grown as ornamental plants in gardens, notably P. setiferum. One species, P. tsus-simense of eastern Asia, is commonly offered as a houseplant.

Related Research Articles

<span class="mw-page-title-main">Thelypteridaceae</span> Family of ferns

Thelypteridaceae is a family of about 900 species of ferns in the order Polypodiales. In the Pteridophyte Phylogeny Group classification of 2016, it is placed in the suborder Aspleniineae. Alternatively, the family may be submerged in a very broadly defined family Aspleniaceae as the subfamily Thelypteridoideae.

<i>Phegopteris</i> Genus of ferns

Phegopteris is a genus of ferns in the family Thelypteridaceae, subfamily Phegopteridoideae, in the Pteridophyte Phylogeny Group classification of 2016. They are known collectively as the beech ferns. Species are native to Asia, North America and Europe.

<i>Pteris</i> Genus of ferns in the subfamily Pteridoideae

Pteris (brake) is a genus of about 300 species of ferns in the subfamily Pteridoideae of the family Pteridaceae. They are native to tropical and subtropical regions, southward to New Zealand, Australia, and South Africa, north to Japan and North America. 78 species are found in China. Some species of Pteris have considerable economic and ecological value, such as Pteris multifida, Pteris ensiformis, Pteris vittata can be used for ornamental purposes; as a hyperaccumulator, Pteris multifida and Pteris vittata can be used to control soil pollution.

<span class="mw-page-title-main">Dryopteridaceae</span> Family of ferns

The Dryopteridaceae are a family of leptosporangiate ferns in the order Polypodiales. They are known colloquially as the wood ferns. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family is placed in the suborder Polypodiineae. Alternatively, it may be treated as the subfamily Dryopteridoideae of a very broadly defined family Polypodiaceae sensu lato.

<i>Cyrtomium</i> Genus of ferns

Cyrtomium is a genus of about 35 species of ferns in the family Dryopteridaceae, subfamily Dryopteridoideae, according to the Pteridophyte Phylogeny Group classification of 2016. Species are native to Asia, Africa, and the Pacific Ocean islands (Hawaii). It is very closely related to the genus Polystichum, with 2016 research suggesting it should be included in a clade sister to Polystichum s.s.

<i>Microsorum</i> Genus of ferns

Microsorum is a genus of ferns in the family Polypodiaceae, subfamily Microsoroideae, according to the Pteridophyte Phylogeny Group classification of 2016 (PPG I). The species are tropical. Like most ferns, they grow from rhizomes, rather than roots. The genus name is often misspelled "Microsorium" or "Microsoreum". It includes some species that are lithophytic rheophytes.

<i>Alsophila</i> (plant) Genus of ferns

Alsophila is a genus of tree ferns in the family Cyatheaceae. It has also been considered to be a section in the subgenus Cyathea of the genus Cyathea.

<i>Aleuritopteris</i> Genus of ferns

Aleuritopteris is a genus of ferns in the Cheilanthoideae subfamily of the Pteridaceae. As with some other genera of the Cheilanthoideae, molecular phylogenetic studies have suggested that it is not monophyletic, and so may need to be circumscribed differently in future.

<i>Arachniodes</i> Genus of ferns

Arachniodes is a fern genus in the family Dryopteridaceae, subfamily Dryopteridoideae, in the Pteridophyte Phylogeny Group classification of 2016. A number of species in this genus are known as "holly ferns".

<i>Bolbitis</i> Genus of ferns

Bolbitis is a genus of ferns in the family Dryopteridaceae, subfamily Elaphoglossoideae, in the Pteridophyte Phylogeny Group classification of 2016.

<i>Tectaria</i> Genus of ferns

Tectaria is a genus of fern in the family Tectariaceae, according to the Pteridophyte Phylogeny Group classification of 2016. Halberd fern is a common name for species in this genus.

<i>Woodsia</i> Genus of ferns

Woodsia is a genus of ferns in the family Woodsiaceae. Species of Woodsia are commonly known as cliff ferns. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), it was the only genus in the family Woodsiaceae. In 2020, Physematium was split off from Woodsia on the basis of molecular phylogenetic evidence. As of June 2023, Plants of the World Online continued to treat Physematium as a synonym of Woodsia.

<i>Deparia</i> Genus of ferns

Deparia is a genus of ferns. The Pteridophyte Phylogeny Group classification of 2016 places the genus in the family Athyriaceae, although other sources include it within an expanded Aspleniaceae or Woodsiaceae.

<i>Christella</i> Genus of ferns

Christella is a genus of around 70-80 species of ferns in the subfamily Thelypteridoideae of the family Thelypteridaceae in the Pteridophyte Phylogeny Group classification of 2016. Other sources sink Christella into a very broadly defined genus Thelypteris. The genus was named after Konrad H. Christ, a Swiss botanist. The distribution of these plants is mostly in the tropics and sub tropical areas. An Australian example is C. dentata.

<i>Pyrrosia</i> Genus of ferns

Pyrrosia is a genus of about 100 fern species in the polypod family, Polypodiaceae. Like other species in Polypodiaceae, the species of Pyrrosia are generally epiphytic on trees or rocks, a few species are terrestrial. The Latin name of Pyrrosia comes from the Greek pyrrhos (red), which refers to its leaves that are red due to the sporangia.

<i>Polystichum mohrioides</i> Species of fern

Polystichum mohrioides is a fern in the family Dryopteridaceae. It is found only in the Falkland and South Georgia Islands in the South Atlantic Ocean.

<i>Goniophlebium</i> Genus of ferns

Goniophlebium is a genus of ferns in the family Polypodiaceae, subfamily Microsoroideae, according to the Pteridophyte Phylogeny Group classification of 2016 (PPG I).

<i>Lepisorus</i> Genus of ferns

Lepisorus is a genus of ferns in the family Polypodiaceae, subfamily Microsoroideae, according to the Pteridophyte Phylogeny Group classification of 2016 (PPG I).

<i>Polystichum drepanum</i> Species of plant

Polystichum drepanum, also known as Madeira shield-fern, is a species of fern in the Dryopteridaceae family.

References

  1. 1 2 3 4 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
  2. ↑ Synonyms Tropicos.org. Missouri Botanical Garden. 25 Jan 2012
  3. 1 2 3 4 Le Péchon, Timothée; Zhang, Liang; He, Hai; Zhou, Xin-Mao; Bytebier, Benny; Gao, Xin-Fen; Zhang, Li-Bing (2016-05-01). "A well-sampled phylogenetic analysis of the polystichoid ferns (Dryopteridaceae) suggests a complex biogeographical history involving both boreotropical migrations and recent transoceanic dispersals". Molecular Phylogenetics and Evolution. 98: 324–336. Bibcode:2016MolPE..98..324L. doi:10.1016/j.ympev.2016.02.018. ISSN   1055-7903. PMID   26944012.
  4. 1 2 Little, D. P.; Barrington, D. S. (2003-03-01). "Major evolutionary events in the origin and diversification of the fern genus Polystichum (Dryopteridaceae)". American Journal of Botany. 90 (3): 508–514. doi: 10.3732/ajb.90.3.508 . ISSN   0002-9122. PMID   21659143. S2CID   11137206.
  5. 1 2 "Polystichum Homepage". www.uvm.edu. Retrieved 2019-12-17.
  6. ↑ Hong-Mei Liu, Robert J. Dyer, Zhi-You Guo, Zhen Meng,Jian-Hui Li, Harald Schneider (2012) The Evolutionary Dynamics of Apomixis in Ferns: A Case Study from Polystichoid Ferns Journal of Botany Volume 2012 (2012), Article ID 510478, 11 pages https://dx.doi.org/10.1155/2012/510478
  7. ↑ Little, Damon P.; Barrington, David S. (2003). "Major evolutionary events in the origin and diversification of the fern genus Polystichum (Dryopteridaceae)". American Journal of Botany. 90 (3): 508–514. doi: 10.3732/ajb.90.3.508 . ISSN   1537-2197. PMID   21659143. S2CID   11137206.
  8. ↑ Li, Chun-xiang; Lu, Shu-gang; Barrington, David S. (2008-01-01). "Phylogeny of Chinese Polystichum (Dryopteridaceae) based on chloroplast DNA sequence data (trnL-F and rps4-trnS)". Journal of Plant Research. 121 (1): 19–26. Bibcode:2008JPlR..121...19L. doi:10.1007/s10265-007-0120-1. ISSN   1618-0860. PMID   18000642. S2CID   35788718.
  9. 1 2 Driscoll, H. E.; Barrington, D. S. (2007-08-01). "Origin of Hawaiian Polystichum (Dryopteridaceae) in the context of a world phylogeny". American Journal of Botany. 94 (8): 1413–1424. doi: 10.3732/ajb.94.8.1413 . ISSN   0002-9122. PMID   21636509. S2CID   5811183.
  10. ↑ Li, Chun-xiang; Lu, Shu-gang; Barrington, David S. (January 2008). "Phylogeny of Chinese Polystichum (Dryopteridaceae) based on chloroplast DNA sequence data (trnL-F and rps4-trnS)". Journal of Plant Research. 121 (1): 19–26. Bibcode:2008JPlR..121...19L. doi:10.1007/s10265-007-0120-1. ISSN   0918-9440. PMID   18000642. S2CID   35788718.
  11. ↑ Hassler, Michael & Schmitt, Bernd (January 2020), "Polystichum", Checklist of Ferns and Lycophytes of the World, 8.20, retrieved 2020-02-02
  12. ↑ NRCS. "Polystichum andersonii". PLANTS Database. United States Department of Agriculture (USDA). Retrieved 19 Jan 2012.
  13. ↑ P. bonseyi Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
  14. ↑ NRCS. "Polystichum bonseyi". PLANTS Database. United States Department of Agriculture (USDA). Retrieved 19 Jan 2012.
  15. ↑ NRCS. "Polystichum brachypterum". PLANTS Database. United States Department of Agriculture (USDA). Retrieved 19 Jan 2012.
  16. ↑ NRCS. "Polystichum braunii". PLANTS Database. United States Department of Agriculture (USDA). Retrieved 19 Jan 2012.
  17. ↑ P. brachypterum Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
  18. ↑ P. capillipes Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
  19. ↑ P. echinatum, Tropicos.org. Missouri Botanical Garden., 19 Jan 2012
  20. ↑ P. kiusiuense Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
  21. ↑ NRCS. "Polystichum haleakalense". PLANTS Database. United States Department of Agriculture (USDA). Retrieved 19 Jan 2012.
  22. ↑ Conspectus of Southern African Pteridophyta, J. P. Roux, Nov. 2001. p. 129
  23. ↑ P. mollissimum Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
  24. ↑ P. nakenense Chinese Plant Names www.eFloras.org 20 Dec 2012
  25. ↑ P. neolobatum Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
  26. ↑ P. ordinatum Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
  27. ↑ P. erinaceum Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
  28. ↑ P. brunneum Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
  29. ↑ NRCS. "Polystichum setigerum". PLANTS Database. United States Department of Agriculture (USDA). Retrieved 20 Jan 2012.
  30. ↑ P. lichiangense Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
  31. ↑ P. sinense Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
  32. ↑ P. apicisterile Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
  33. ↑ P. integripinnulum Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
  34. ↑ P. conaense Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
  35. ↑ P. ilicifolium Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
  36. ↑ P. heteropaleaceum Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
  37. ↑ P. kodamae Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
  38. ↑ P. xiphophyllum Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
  39. ↑ P. gyirongense Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
  40. ↑ P. jizhushanense Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
  41. ↑ Murphy, Rosaline J; Page, Christopher N; Parslow, Rosemary E; Bennallick, Ian J (2012). Ferns, Clubmosses, Quillworts and Horsetails of Cornwall and the Isles of Scilly. Truro: ERCCIS. ISBN   978-1-902864-07-5.
  42. ↑ P. auriculatum Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
  43. ↑ P. falcatum Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
  44. ↑ P. lepidocaulon Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
  45. ↑ LI, Chunxiang (2004). "Asian origin for Polystichum (Dryopteridaceae) based on rbcL sequences". Chinese Science Bulletin. 49 (11): 1146. Bibcode:2004ChSBu..49.1146L. doi:10.1360/04wc0086. ISSN   1001-6538. S2CID   85087849.
  46. ↑ Morero, Rita E.; Deanna, Rocío; Barboza, Gloria E.; Barrington, David S. (August 2019). "Historical biogeography of the fern genus Polystichum (Dryopteridaceae) in Austral South America". Molecular Phylogenetics and Evolution. 137: 168–189. Bibcode:2019MolPE.137..168M. doi: 10.1016/j.ympev.2019.05.004 . PMID   31077789. S2CID   150375407.
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