Asplenium scolopendrium

The most striking and unusual feature of the fern is its simple, strap-shaped undivided fronds. The supposed resemblance of the leaves to the tongue of a hart (an archaic term for a male red deer) gave rise to the common name "hart's-tongue fern".

File:Aspleniaceae Asplenium scolopendrium 1.jpg

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File:Asplenium scolopendrium prothallus.jpg

File:Jeune sporophyte d'Asplenium scolopendrium (focus).jpg

Linnaeus first gave the hart's-tongue fern the binomial Asplenium scolopendrium in his Species Plantarum of 1753.{{sfn|Linnaeus|1753|p=1079}} The Latin specific epithet scolopendrium is derived from the Greek skolopendra, meaning a centipede or millipede; this is due to the sori pattern being reminiscent of a myriapod's legs.{{cite book | last=Harrison |first=Lorraine | title=RHS Latin for Gardeners | year=2012 |publisher=Mitchell Beazley | location=United Kingdom | isbn=978-1845337315 }}{{Cite web|url=https://websites.rbge.org.uk/cgi-bin/nph-readbtree.pl/feout?FAMILY_XREF=&GENUS_XREF=Asplenium+&SPECIES_XREF=scolopendrium&TAXON_NAME_XREF=&RANK=|title=Flora Europaea Search Results|website=Websites.rbge.org.uk|access-date=16 March 2022}}

A global phylogeny of Asplenium published in 2020 divided the genus into eleven clades,{{sfn|Xu|Zhang|Rothfels|Smith|2020|p=27}} which were given informal names pending further taxonomic study. A. scolopendrium belongs to the "Phyllitis subclade" of the "Phyllitis clade".{{sfn|Xu|Zhang|Rothfels|Smith|2020|p=30}} Members of the Phyllitis clade have undivided or pinnatifid leaf blades with a thick, leathery texture, persistent scales on their stalk, and often possess anastomosing veins. Members of the Phyllitis subclade have undivided leaves with freely branching veins and single or paired sori. They are widely distributed through the Northern Hemisphere.{{sfn|Xu|Zhang|Rothfels|Smith|2020|p=41}} If defined to include the former A. komarovii, A. scolopendrium makes up the former segregate genus Phyllitis and is sister to A. sagittatum.{{sfn|Xu|Zhang|Rothfels|Smith|2020|p=30}}

Three subspecies were accepted in the most recent revision of the species:{{cite journal |last1=Heo |first1=Namjoo |last2=Yun |first2=Seona |last3=Fernando |first3=Danilo D. |title=Molecular phylogenetic assessment of three major taxa in the Asplenium scolopendrium complex (Aspleniaceae) |journal=Taxon |volume=72 |issue=2 |pages=245–260 |year=2023 |doi=10.1002/tax.12890 |url=https://onlinelibrary.wiley.com/doi/10.1002/tax.12890}}

• A. scolopendrium subsp. scolopendrium is native to Europe, North Africa, the Middle East, and Central Asia. Populations in the eastern Mediterranean have been referred to as A. scolopendrium var. antri-jovis.
• A. scolopendrium subsp. americanum is native to Ontario in Canada, the United States, southern Mexico, and Hispaniola. The tropical populations have been referred to as A. scolopendrium var. lindenii.
• A. scolopendrium subsp. japonicum, formerly A. komarovii, is native to eastern Russia, China, Taiwan, and Japan

Morphological differences between the varieties are minor, but the North American variety americanum is tetraploid, while the Old World subspecies A. scolopendrium scolopendrium) is diploid.{{Cite web|url=http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500199|title=Asplenium scolopendrium var. americanum in Flora of North America|website=Efloras.org|access-date=16 March 2022}}

Asplenium scolopendrium is a common species in the Old World:

• The subspecies scolopendrium occurs throughout Europe (including the Caucasus and the British Isles). It is absent however in northeast Europe (Finland, Belarus, Latvia, Lithuania and European Russia).Christenhusz, M., Bento Elias, R., Dyer, R., Ivanenko, Y., Rouhan, G., Rumsey, F. & Väre, H. 2017. Asplenium scolopendrium (Europe assessment). The IUCN Red List of Threatened Species 2017: e.T83504456A83506952. Accessed on 19 June 2024.{{Cite journal |last1=Heo |first1=Namjoo |last2=Lomolino |first2=Mark V |last3=Watkins |first3=James E |last4=Yun |first4=Seona |last5=Weber-Townsend |first5=Josh |last6=Fernando |first6=Danilo D |date=2022-09-01 |title=Evolutionary history of the Asplenium scolopendrium complex (Aspleniaceae), a relictual fern with a northern pan-temperate disjunct distribution |url=https://doi.org/10.1093/biolinnean/blac080 |journal=Biological Journal of the Linnean Society |volume=137 |issue=2 |pages=183–199 |doi=10.1093/biolinnean/blac080 |issn=0024-4066}} Beyond Europe, it is found in the Middle East, and North Africa (in Morocco, Algeria, Tunisia and Libya, as well as the Canary Islands). It has also been introduced to the Falkland Islands. Unlike its American counterpart, the scolopendrium of Europe is used for many horticultural uses.{{Cite journal |last=Pence |first=Valerie C. |date=2015 |title=Propagation and Cryopreservation of Asplenium scolopendrium var. americanum, the American Hart's-Tongue Fern |url=https://www.jstor.org/stable/44076071 |journal=American Fern Journal |volume=105 |issue=3 |pages=211–225 |issn=0002-8444}}{{Cite web|url=http://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:30216020-2|title=Asplenium scolopendrium L. {{pipe}} Plants of the World Online {{pipe}} Kew Science|website=Plants of the World Online|access-date=16 March 2022}} Specimens of this variety found in North America (such as in New Brunswick and Ontario in Canada, and Maryland in the United States) are considered naturalized descendants of cultivated plants.Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland. First edition. In: Kartesz, J.T., and C.A. Meacham. Synthesis of the North American Flora, Version 1.0. North Carolina Botanical Garden, Chapel Hill, N.C.

In North America, it occurs in rare, widely scattered populations located in different locales:

• in the Mexican states of Chiapas, Coahuila, Nuevo León, and Oaxaca, as well as the Caribbean island of Hispaniola (in Haiti).{{Cite web|url=http://legacy.tropicos.org/Name/26603800?tab=distribution|title=Tropicos {{pipe}} Name - Asplenium scolopendrium L.|website=Legacy.tropicos.org|access-date=16 March 2022}}
• along the Onondaga Limestone and Niagara Escarpment geological formations in Central New York (present in 2 counties), southern Ontario (present in 7 counties), and the eastern Upper Peninsula in Michigan (present in 2 counties). Exceptions are disjunct populations that exist in Alabama (in Fern Cave National Wildlife Refuge, a wildlife refuge centered around an off-limits cave in Jackson County, Alabama, where it has declined heavily due to illegal plant collecting, and an undisclosed pit in Morgan County that is also off-limits and protected), and Tennessee (in just a single county); these southern populations are at dire risk of extirpation.{{cite web|url=https://ecos.fws.gov/ServCat/DownloadFile/174090|title=Species Status Assessment : Report for the American Hart's-tongue Fern (Asplenium scolopendrium var. americanum)|date=November 2019|website=Ecos.fws.gov|access-date=16 March 2022}} An introduced population descended from New York plants is found in New Jersey; it is a remnant of a 1936 effort to practice ex-situ conservation of populations in New York.Snyder, D.B. 1990. Botanist, New Jersey Natural Heritage Program. Personal communication with Wayne Ostlie, MRO, The Nature Conservancy.
• In 2020, a new population of hart's-tongue ferns was discovered inside a cave with basaltic lava flows in El Malpais National Monument, Cibola County, New Mexico; this represents the first confirmed population of the species in North America west of the Mississippi. Genetic analyses and surveys are currently being performed to determine the population's variation and overall health. {{Cite web |url=https://www.fws.gov/northeast/EcologicalServices/pdf/endangered/20200729_5YR_AmericanHarts_tongue_fern.pdf |title=Archived copy |access-date=2021-11-11 |archive-date=2021-11-11 |archive-url=https://web.archive.org/web/20211111092007/https://www.fws.gov/northeast/EcologicalServices/pdf/endangered/20200729_5YR_AmericanHarts_tongue_fern.pdf |url-status=dead }}

In East Asia, A. scolopendrium subsp. japonicum is distributed in the Russian Far East, north-eastern China and the Korean Peninsula, however it is considered as being generally rare on the mainland.Lee, Y. M., and W. Y. Lee. "Illustrated rare and endangered species in Korea." Korea National Arboretum. Po-cheon, Korea (1997): 41.Lin YX, Viane R. 2013. Aspleniaceae. In: Wu ZY, Raven PH, Hong DY, eds. Flora of China, Vol. 2-3. Beijing: Science Press; St. Louis: Missouri Botanical Garden Press, 274.Dong, Shiyong, et al. "Red list assessment of lycophytes and ferns in China." Biodiversity Science 25.7 (2017): 765. In contrast, it is relatively abundant on many islands including the Japanese Archipelago.

The unique dispersal of Asplenium scolopendrium has caught the attention of international botanists. In fact, the very existence of such varieties beg that "...these populations arose following colonization events involving a single spore".Weston L. Testo, James E. Watkins Jr, John Wiley, Laura Baumann, Eric Weaver "SHORTER NOTE," American Fern Journal, 111(3), 217-222, (2 September 2021). Retrieved March 17, 2023, from https://bioone.org/journals/american-fern-journal/volume-111/issue-3/0002-8444-111.3.217/SHORTER-NOTE/10.1640/0002-8444-111.3.217.short

The plants grow on neutral, calcium-rich, and/or lime-rich substrates under deciduous hardwood canopies (usually sugar maples in eastern North America), including moist soil and damp crevices in old walls; they are found most commonly in shaded areas. Plants in full sun are usually stunted and yellowish in colour, while those in full shade are dark green and healthy. The disjunct populations of the North American variation in the southeastern US are found exclusively in sinkhole pits or limestone caves. These populations may be relics of cooler Pleistocene climates.{{cite book|last1=Short|first1=John W.|last2=Spaulding|first2=Daniel D.|title=Ferns of Alabama|date=2012|publisher=University of Alabama Press|location=Tuscaloosa, AL|isbn=9780817356477}}

The most recent International Union for Conservation of Nature (IUCN) assessment for The IUCN Red List of Threatened Species was conducted in 2016, with the conservation status of Asplenium scolopendrium being listed as being a species of Least Concern (LC). Despite the positive status, this assessment is chiefly limited to the conservation situation of A. scolopendrium in Europe, as opposed to other parts of its natural range.

Asplenium scolopendrium is found across the vast majority of European countries, in a wide variety of natural settings, across a multitude of habitats. It is also able to colonise manmade environments including growing in walls and brickwork crevices, particularly in limestone mortar. Furthermore, it is considered to be a gregarious species - it is able to spread both sexually and asexually, frequently forming large clumps and present in high densities - in some areas it is considered very common.

Across Europe as a whole, the IUCN assessed no immediate threats to this species in natural settings, and predicted that the species may benefit from milder wetter winters predicted under various climate change scenarios. In the urban environment, where A. scolopendrium frequently grows on vertical faces and in crevices of lime mortared brick walls, the species may be threatened by wall renovation projects. Although the fern is often cultivated as a garden plant or used in landscape planting, commercially sold plants are sourced from horticultural stock.{{Cite web |title=Asplenium scolopendrium, hart's tongue fern |url=http://www.rhs.org.uk/plants/25910/asplenium-scolopendrium/details |access-date=2024-06-27 |website=www.rhs.org.uk |language=en-gb}}

Asplenium scolopendrium is not a specifically protected species in the majority of European states, however in part due to its wide distribution, A. scolopendrium inhabits a large number of legally protected areas across the continent. Despite this, it is considered to be declining in numbers or a species at risk in certain parts of its range. Asplenium scolopendrium was first listed as "Vulnerable" in the National Red Lists for Albania in 2014{{Cite web|url=https://www.nationalredlist.org/|title=The National Red List Project |website=Nationalredlist.org |access-date=16 March 2022}} and Norway in 2010 (under Criterion D1, that is, having a very small population of under 1000 individuals).Kålås, J.A., Viken, Å. and Bakken, T. (eds). 2006. Norsk Rødliste 2006 – 2006 Norwegian Red List. Artsdatabanken. It is considered "critically threatened and rare" in the Czech Republic's 2012 plant Red List;Grulich, V. 2012. Red List of vascular plants of the Czech Republic: 3rd edition. Preslia 84: 631-645. and "Endangered" in Sweden's 2010 Red List.Gärdenfors, U. 2010. Rödlistade arter i Sverige - The 2010 Red List of Swedish Species. ArtDatabanken, SLU, Uppsala.

In the United States, A. scolopendrium var. americanum was declared endangered by the U.S. Fish and Wildlife Service in 1989.{{cite report |title=American hart's-tongue recovery plan |last=Currie |first=Robert R. |date=September 1993 |url=http://www.fws.gov/northeast/nyfo/es/amhtfrecovplan.pdf |publisher=U.S. Fish and Wildlife Service |location=Atlanta, Georgia |access-date=2010-02-28}} The reasons for its rarity are currently being researched, with reintroduction programs in New York and elsewhere also in development.{{cite web|url=https://digitalcommons.esf.edu/cgi/viewcontent.cgi?article=1008&context=etds|title=Experimental Reintroduction of American Hart's-Tongue Fern (Asplenium scolopendrium var. americanum) : Factors Affecting Successful Establishment of Transplants|author=Michael Serviss|website=Digitalcommons.esf.edu|access-date=16 March 2022}}

Ontario, Canada has the highest population numbers of A. scolopendrium var. americanum of any region in the variation's distribution, with around 80% of all subpopulations and around 94% of all individuals. The fern was reported at more than 100 sites across the province, with around 75 still believed to be existing. Despite this, A. scolopendrium var. americanum was listed as a species of Special Concern under the Committee on the Status of Species at Risk in Ontario in May 2017, due to its extremely specific habitat requirements, relatively small distribution, and some subpopulations consisting of too little individuals.{{cite web|url=http://cossaroagency.ca/wp-content/uploads/2018/06/AccessibleCOSSAROEvaluation_American_Harts-tongue_Fern_FINAL_30JAN2018.pdf|title=Ontario Species at Risk Evaluation Report for American Hart's-tongue Fern (Asplenium scolopendrium var. americanum)|website=Cossaroagency.ca|access-date=16 March 2022}}{{Cite web|url=http://www.ontario.ca/page/american-harts-tongue-fern|title=American Hart's Tongue Fern|website=Ontario.ca|access-date=16 March 2022}}

Asplenium scolopendrium is often grown as an ornamental plant, with several cultivars selected with varying frond form, including with frilled frond margins, forked fronds and cristate forms. The species has gained the Royal Horticultural Society's Award of Garden Merit,{{cite web |title=RHS Plant Selector - Asplenium scolopendrium |url=https://www.rhs.org.uk/Plants/25910/Asplenium-scolopendrium/Details |website=Rhs.org.uk|access-date=12 February 2020}} as has the cultivar

'Angustatum'.{{cite web | url = https://www.rhs.org.uk/Plants/48025/Asplenium-scolopendrium-Angustatum/Details

| title = Asplenium scolopendrium 'Angustatum'|website=Rhs.org.uk|access-date = 20 February 2020}}

The American variety is reputed to be difficult to cultivate (making conservation efforts for it even more troublesome); due to this, most, if not all, cultivated individuals are derived from the Old World subspecies.{{cite book |title=Ferns for American Gardens |first=John T. |last=Mickel |year=2003 |publisher=Timber Press |isbn=9780881925982}} This book is a reprinting of {{cite book |title=Ferns for American Gardens |url=https://archive.org/details/isbn_9780025844919 |url-access=registration |first=John T. |last=Mickel |publisher=MacMillan |year=1994|isbn=9780025844919 }}

This fern was used in the 1800s as a medicinal plant in folk medicine as a spleen tonic (hence an archaic name for the genus, "spleenworts") and for other uses.{{cite book |title=The family herbal: or An account of all those English plants, which are remarkable for their virtues, and of the drugs which are produced by vegetables of other countries; with their descriptions and their uses, as proved by experience |last=Hill |first=John |year=1812 |publisher=C. Brightly and T. Kinnersley |page=[https://archive.org/details/familyherbaland00kinngoog/page/n261 162] |url=https://archive.org/details/familyherbaland00kinngoog}}

A manuscript written by Katharine Palmer in the 18th century contains a recipe for hart's-tongue ale.{{cite book |title=A collection of ye best receipts most approved and fittest in cookery, preserving, and all manner of housewifery, physick & chirurgery |url=https://wellcomecollection.org/works/mde4asuj |author=Palmer, Katharine |year=1700 |page=[https://wellcomecollection.org/works/mde4asuj/items?canvas=80 77]|publisher=MS.7976 (wellcome collection)}}

Phyllitis scolopendrium habitat.jpg|Habitat

Phyllitis scolopendrium3.jpg|{{center|A cultivar with frilled frond margins}}

Spiers harts tongue fern.JPG|{{center|A specimen growing in lime mortar on a wall}}

Asplenium scolopendrium americanum.jpg|{{center|Example of the North American variety}}

{{Reflist}}

• {{cite book | title=Species Plantarum | edition=1st | volume=II | last=Linnaeus | first=C. | authorlink=Carl Linnaeus | year=1753 | publisher=Laurentii Salvii | location=Stockholm | url=https://www.biodiversitylibrary.org/page/359100}}
• {{cite journal |last1=Xu |first1=Ke-Wang |last2=Zhang |first2=Liang |last3=Rothfels |first3=Carl J. |last4=Smith |first4=Alan R. |last5=Viane |first5=Ronald |last6=Lorence |first6=David |last7=Wood |first7=Kenneth R. |last8=Cheng |first8=Cheng-Wei |last9=Knapp |first9=Ralf |last10=Zhou |first10=Lin |last11=Lu |first11=Ngan Thi |last12=Zhou |first12=Xin-Mao |last13=Wei |first13=Hong-Jin |last14=Fan |first14=Qiang |last15=Chen |first15=Su-Fang |last16=Cicuzza |first16=Daniele |last17=Gao |first17=Xin-Fen |last18=Li |first18=Wen-Bo |last19=Zhang |first19=Li-Bing |title=A global plastid phylogeny of the fern genus Asplenium (Aspleniaceae) |year=2020 |journal=Cladistics |volume=36 |issue=1 |pages=22–71 |doi=10.1111/cla.12384|pmid=34618950 |s2cid=201197385 |doi-access=free }}

• Hyde, H. A., Wade, A. E., & Harrison, S. G. (1978). Welsh Ferns. National Museum of Wales. {{ISBN|0-7200-0210-9}}.
• {{cite web |title=A Real Rarity |publisher=Finger Lakes Native Plant Society |url=http://flnps.org/native-plants/real-rarity |date=December 2009 |last=Parker |first=Rosemarie}} A popular article on hart's tongue fern that includes several references and a discussion of cultivation possibilities for the European and American varieties. The article strongly discourages collection and or cultivation of the North American variety.

{{Commons category|Asplenium scolopendrium|Asplenium scolopendrium}}

• [https://www.youtube.com/watch?v=aVaIhg3krCI Commentary and video on Hart's Tongue ferns at Lynn Glen, Dalry, Scotland]

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scolopendrium

Category:Garden plants of North America

Category:Medicinal plants of North America

Category:Plants described in 1753

Category:Ferns of Europe

Category:Ferns of Asia

Category:Ferns of the Americas

Category:Taxa named by Carl Linnaeus