Polystichum is a genus of ferns in the familyDryopteridaceae, 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]
Description
Many ferns of this genus have stout, slowly creeping rootstocks that form a crown, with a vase-like ring of evergreenfronds 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[update], 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.
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]
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.
References
^ abcdPPG 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, S2CID39980610
^Synonyms Tropicos.org. Missouri Botanical Garden. 25 Jan 2012
^ abcdLe 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. ISSN1055-7903. PMID26944012.
^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
^P. kodamae Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
^P. xiphophyllum Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
^P. gyirongense Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
^P. jizhushanense Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
^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. ISBN978-1-902864-07-5.
^P. auriculatum Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
^P. falcatum Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
^P. lepidocaulon Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
