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Polystichum

( Dryopteridaceae )

Description

Apomixis

Taxonomy

Selected species
Hybrids
Former species

Distribution
Ecology
Cultivation
References
External links

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Fern Polystichum Species Tropicos

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Polystichum is a genus of in the family Dryopteridaceae, subfamily Dryopteridoideae, according to the Pteridophyte Phylogeny Group classification of 2016 (PPG I). The genus has about 500 species and has a cosmopolitan distribution. The highest diversity is in eastern Asia, with about 208 species in China alone; 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, scree slopes, and stream banks.

Description

Many ferns of this genus have stout, slowly creeping rootstocks that form a crown, with a vase-like ring of evergreen long. The Sorus are round, with a circular indusium, except in South American species which lack an indusium. The stipes have prominent scales and often have hair-like cilia, but they lack any true hairs. 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

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Taxonomy

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

Selected species

The genus has a large number of species. The PPG I classification suggested that there were about 500 species; , the Checklist of Ferns and Lycophytes of the World listed 397 species and 58 hybrids, noting that "many undescribed species remain". 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.

– Christmas fern
– Hard shield fern
(=)
– Aleutian shield fern
– Anderson's hollyfern
Polystichum atkinsonii
Polystichum australiense
Polystichum bakerianum
Polystichum biaristatum
Polystichum bomiense
Polystichum bonapartii
Polystichum bonseyi P. bonseyi Tropicos.org. Missouri Botanical Garden. 19 Jan 2012 – Bonsey's Hollyfern
Polystichum brachypterum – Rusty swordfern
Polystichum braunii – Braun's shield fern, Braun's hollyfern
Polystichum brachypterum P. brachypterum Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
(=) Polystichum garhwalicum
Polystichum bulbiferum
Polystichum calderonense – Monte Guilarte hollyfern
Polystichum californicum – California sword fern
Polystichum capillipes P. capillipes Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
(=) Polystichum michelii
(=) Polystichum minusculum
(=) Polystichum molliculum
Polystichum castaneum
Polystichum chilense
Polystichum christii
Polystichum chunii
Polystichum craspedosorum
Polystichum cyclolobum
Polystichum cystostegia – Alpine shield fern
Polystichum deltodon
Polystichum dielsii
Polystichum discretum
Polystichum drepanum
Polystichum dudleyi – Dudley's sword fern
Polystichum duthiei
(=) Polystichum tsuchuense
Polystichum echinatum – Rusty swordfern
Polystichum erosum
Polystichum excellens
Polystichum eximium
(=) Polystichum tialooshanense
Polystichum falcatipinnum
Polystichum falcinellum
Polystichum fallax
Polystichum formosanum
Polystichum glandulosum
Polystichum gongboense
(=) Polystichum rarum
Polystichum grandifrons
(=) Polystichum kiusiuense P. kiusiuense Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
Polystichum gymnocarpium
Polystichum haleakalense – Island swordfern
Polystichum hancockii
Polystichum hecatopteron
Polystichum herbaceum
Polystichum imbricans – Narrowleaf sword fern
Polystichum incongruum
Polystichum kruckebergii – Kruckeberg's sword fern/holly fern
Polystichum kwakiutlii
Polystichum lachenense
Polystichum lanceolatum
Polystichum lemmonii – Lemmon's holly fern, Shasta fern
Polystichum lentum
Polystichum lonchitis – Holly fern
Polystichum longidens
Polystichum longipaleatum
Polystichum longipes
Polystichum luctuosum – Mourning shield fern
Polystichum macleae Conspectus of Southern African Pteridophyta, J. P. Roux, Nov. 2001. p. 129
Polystichum macrochlaenum
Polystichum makinoi
Polystichum martini
Polystichum mayebarae
Polystichum mediocre
Polystichum medogense
Polystichum microchlamys
Polystichum mohrioides
Polystichum mollissimum P. mollissimum Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
Polystichum monticola – Mountain shield fern
Polystichum moorei
Polystichum morii
Polystichum moupinense
(=) Polystichum woodsioides
Polystichum munitum – Western sword fern
(=) Polystichum solitarium
Polystichum muricatum
(=) Polystichum moritzianum
Polystichum nakenense P. nakenense Chinese Plant Names www.eFloras.org 20 Dec 2012
Polystichum neolobatum P. neolobatum Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
(=) Polystichum assurgens
(=) Polystichum yigongense
Polystichum nepalense
(=) Polystichum atroviridissimum
Polystichum ningshenense
Polystichum obliquum
Polystichum omeiense
(=) Polystichum carvifolium
Polystichum ordinatum
(=) Polystichum bicolor P. ordinatum Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
Polystichum orientalitibeticum
Polystichum paramoupinense
Polystichum parvipinnulum
Polystichum piceopaleaceum
Polystichum polyblepharum – Tassel fern
(=) Polystichum aculeatum var. japonicum
(=) Polystichum setosum
Polystichum prescottianum
(=) Polystichum erinaceum P. erinaceum Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
Polystichum prionolepis
(=) Polystichum rectipinnum
Polystichum proliferum – Mother shield fern
Polystichum pseudocastaneum
(=) Polystichum brunneum P. brunneum Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
Polystichum pseudomakinoi
Polystichum punctiferum
(=) Polystichum glingense
(=) Polystichum virescens
Polystichum pungens – Forest shield fern
(=) Polystichum lucidum
Polystichum qamdoense
(=) Polystichum tumbatzense
Polystichum retrosopaleaceum
(=) Polystichum aculeatum var. retrosopaleaceum
Polystichum rhombiforme
Polystichum rhomboidea
(=) Polystichum rhomboideum
Polystichum richardii – Common shield fern
Polystichum rigens
(=) Polystichum platychlamys
Polystichum rotundilobum
Polystichum scopulinum – Mountain holly fern, rock sword fern
(=) Polystichum mohrioides var. scopulinum
Polystichum semifertile
(=) Polystichum nyalamense
Polystichum setiferum – Soft shield fern
(=) Polystichum angulare
Polystichum setigerum – Alaska hollyfern
Polystichum shensiense
(=) Polystichum lichiangense P. lichiangense Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
Polystichum silvaticum
Polystichum simplicipinnum
Polystichum sinense P. sinense Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
(=) Polystichum atuntzeense
(=) Polystichum austrotibeticum
(=) Polystichum decorum
(=) Polystichum ellipticum
(=) Polystichum fukuyamae
(=) Polystichum lhasaense
(=) Polystichum parasinense
(=) Polystichum wilsoni
Polystichum squarrosum
(=) Polystichum apicisterile P. apicisterile Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
(=) Polystichum integripinnulum P. integripinnulum Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
Polystichum stenophyllum
var. conaense (=) Polystichum conaense P. conaense Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
Polystichum stimulans
(=) Polystichum ilicifolium P. ilicifolium Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
Polystichum submite
(=) Polystichum paradoxum
Polystichum tacticopterum
(=) Polystichum heteropaleaceum P. heteropaleaceum Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
(=) Polystichum kodamae P. kodamae Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
Polystichum thomsoni
Polystichum tibeticum
Polystichum transvaalense – Stemmed shield fern
Polystichum tripteron
Polystichum tsus-simense – Korean rock fern
(=) Polystichum falcilobum
Polystichum vestitum – Prickly shield fern
Polystichum wattii
Polystichum whiteleggei
Polystichum xiphophyllum P. xiphophyllum Tropicos.org. Missouri Botanical Garden. 20 Jan 2012
(=) Polystichum monotis
(=) Polystichum praelongum
Polystichum yadongense
Polystichum yunnanense
(=) Polystichum gyirongense P. gyirongense Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
(=) Polystichum jizhushanense P. jizhushanense Tropicos.org. Missouri Botanical Garden. 20 Jan 2012

Hybrids

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

P. × bicknellii ( P. aculeatum × P. setiferum)
P. × illyricum ( P. aculeatum × P. lonchitis)
P. × lonchitiforme ( P. lonchitis × P. setiferum)
P. × lesliei ( P. setiferum × P. munitum) first found in Surrey in 1995 and a second plant found in Cornwall in 2001.
Rosaline J Murphy (2025). 9781902864075, ERCCIS. ISBN 9781902864075
P. × potteri ( P. braunii × P. acrostichoides)

Former species

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

Polystichum auriculatum = Dryopteris auriculata P. auriculatum Tropicos.org. Missouri Botanical Garden. 19 Jan 2012
Polystichum falcatum = Cyrtomium falcatum P. falcatum Tropicos.org. Missouri Botanical Garden. 19 Jan 2012 – Japanese Holly Fern
Polystichum lepidocaulon = Cyrtomidictyum lepidocaulon P. lepidocaulon Tropicos.org. Missouri Botanical Garden. 20 Jan 2012

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. During this period there were high temperatures across the globe, which may have contributed to the diversification of flora.

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 Beringia. 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.

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

Ecology

Polystichum species are used as food plants by the 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 in gardens, notably P. setiferum. One species, P. tsus-simense of eastern Asia, is commonly offered as a houseplant.

External links