Hart's-tongue
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''Asplenium scolopendrium'', commonly known as the hart's-tongue fern, is an
evergreen In botany, an evergreen is a plant which has foliage that remains green and functional through more than one growing season. This also pertains to plants that retain their foliage only in warm climates, and contrasts with deciduous plants, which ...
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except t ...
in the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
'' Asplenium'' native to the Northern Hemisphere.


Description

The most striking and unusual feature of the fern is its simple, undivided fronds. The leaves' supposed resemblance to the tongue of a hart (an archaic term for a male
red deer The red deer (''Cervus elaphus'') is one of the largest deer species. A male red deer is called a stag or hart, and a female is called a hind. The red deer inhabits most of Europe, the Caucasus Mountains region, Anatolia, Iran, and parts of wes ...
) gave rise to the common name "hart's-tongue fern".


Taxonomy

Linnaeus first gave the hart's-tongue fern the
binomial Binomial may refer to: In mathematics *Binomial (polynomial), a polynomial with two terms * Binomial coefficient, numbers appearing in the expansions of powers of binomials *Binomial QMF, a perfect-reconstruction orthogonal wavelet decomposition ...
''Asplenium scolopendrium'' in his ''
Species Plantarum ' (Latin for "The Species of Plants") is a book by Carl Linnaeus, originally published in 1753, which lists every species of plant known at the time, classified into genera. It is the first work to consistently apply binomial names and was the ...
'' of 1753. The Latin
specific epithet In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
''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. A global phylogeny of ''Asplenium'' published in 2020 divided the genus into eleven clades, which were given informal names pending further taxonomic study. ''A. scolopendrium'' belongs to the "''Phyllitis'' subclade" of the "''Phyllitis'' clade". 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. Its closest relative within the subclade is '' A. komarovii'', with which it forms a clade (the former segregate genus ''Phyllitis''), sister to '' A. sagittatum''. Three variations or subspecies are currently known: *''A. scolopendrium'' var. ''scolopendrium'', the most common variation, is native to
Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelago a ...
and
North Africa North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...
; *''A. scolopendrium'' var. ''americanum'', a threatened variation, is native to the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
and
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...
; *''A. scolopendrium'' var. ''lindenii'', the least-studied variation, is native to
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
and
Hispaniola Hispaniola (, also ; es, La Española; Latin and french: Hispaniola; ht, Ispayola; tnq, Ayiti or Quisqueya) is an island in the Caribbean that is part of the Greater Antilles. Hispaniola is the most populous island in the West Indies, and th ...
. Morphological differences between the variations are minor, but the North American variations (i.e. ''americanum'' and ''lindenii'') are tetraploid, while the
Old World The "Old World" is a term for Afro-Eurasia that originated in Europe , after Europeans became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia, which were previously thought of by the ...
variation (i.e. ''scolopendrium'') is
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
. Some taxonomists group ''lindenii'' with ''americanum'', leaving just two variations.


Distribution

''Asplenium scolopendrium'' is a common species in the Old World: *The variation ''scolopendrium'' has been found throughout
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
(including the
Caucasus The Caucasus () or Caucasia (), is a region between the Black Sea and the Caspian Sea, mainly comprising Armenia, Azerbaijan, Georgia, and parts of Southern Russia. The Caucasus Mountains, including the Greater Caucasus range, have historically ...
and the
British Isles The British Isles are a group of islands in the North Atlantic Ocean off the north-western coast of continental Europe, consisting of the islands of Great Britain, Ireland, the Isle of Man, the Inner and Outer Hebrides, the Northern Isles, ...
), the
Middle East The Middle East ( ar, الشرق الأوسط, ISO 233: ) is a geopolitical region commonly encompassing Arabian Peninsula, Arabia (including the Arabian Peninsula and Bahrain), Anatolia, Asia Minor (Asian part of Turkey except Hatay Pro ...
, and North Africa (in
Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
,
Algeria ) , image_map = Algeria (centered orthographic projection).svg , map_caption = , image_map2 = , capital = Algiers , coordinates = , largest_city = capital , relig ...
,
Tunisia ) , image_map = Tunisia location (orthographic projection).svg , map_caption = Location of Tunisia in northern Africa , image_map2 = , capital = Tunis , largest_city = capital , ...
and
Libya Libya (; ar, ليبيا, Lībiyā), officially the State of Libya ( ar, دولة ليبيا, Dawlat Lībiyā), is a country in the Maghreb region in North Africa. It is bordered by the Mediterranean Sea to the north, Egypt to Egypt–Libya bo ...
, as well as the
Canary Islands The Canary Islands (; es, Canarias, ), also known informally as the Canaries, are a Spanish autonomous community and archipelago in the Atlantic Ocean, in Macaronesia. At their closest point to the African mainland, they are west of Morocc ...
); it has also been introduced to the
Falkland Islands The Falkland Islands (; es, Islas Malvinas, link=no ) is an archipelago in the South Atlantic Ocean on the Patagonian Shelf. The principal islands are about east of South America's southern Patagonian coast and about from Cape Dubouzet ...
of
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
. Specimens of this variation found in North America (such as in
New Brunswick New Brunswick (french: Nouveau-Brunswick, , locally ) is one of the thirteen provinces and territories of Canada. It is one of the three Maritime provinces and one of the four Atlantic provinces. It is the only province with both English and ...
and
Ontario Ontario ( ; ) is one of the thirteen provinces and territories of Canada.Ontario is located in the geographic eastern half of Canada, but it has historically and politically been considered to be part of Central Canada. Located in Central Ca ...
in Canada, and
Maryland Maryland ( ) is a state in the Mid-Atlantic region of the United States. It shares borders with Virginia, West Virginia, and the District of Columbia to its south and west; Pennsylvania to its north; and Delaware and the Atlantic Ocean to ...
in the United States) are considered naturalized descendants of cultivated plants. In
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
, it occurs in rare, widely scattered populations located in different locales: *The variation ''lindenii'' grows in the Mexican states of
Chiapas Chiapas (; Tzotzil language, Tzotzil and Tzeltal language, Tzeltal: ''Chyapas'' ), officially the Free and Sovereign State of Chiapas ( es, Estado Libre y Soberano de Chiapas), is one of the states that make up the Political divisions of Mexico, ...
,
Coahuila Coahuila (), formally Coahuila de Zaragoza (), officially the Free and Sovereign State of Coahuila de Zaragoza ( es, Estado Libre y Soberano de Coahuila de Zaragoza), is one of the 32 states of Mexico. Coahuila borders the Mexican states of N ...
,
Nuevo León Nuevo León () is a state in the northeast region of Mexico. The state was named after the New Kingdom of León, an administrative territory from the Viceroyalty of New Spain, itself was named after the historic Spanish Kingdom of León. With a ...
, and
Oaxaca Oaxaca ( , also , , from nci, Huāxyacac ), officially the Free and Sovereign State of Oaxaca ( es, Estado Libre y Soberano de Oaxaca), is one of the 32 states that compose the political divisions of Mexico, Federative Entities of Mexico. It is ...
, as well as the
Caribbean The Caribbean (, ) ( es, El Caribe; french: la Caraïbe; ht, Karayib; nl, De Caraïben) is a region of the Americas that consists of the Caribbean Sea, its islands (some surrounded by the Caribbean Sea and some bordering both the Caribbean Se ...
island of Hispaniola. *The variation ''americanum'' mostly grows along the
Onondaga Limestone The Onondaga Limestone is a group of hard limestones and dolomites of Devonian age that form an important geographic feature in some areas in which it outcrops; in others, especially its Southern Ontario portion, the formation can be less promine ...
and
Niagara Escarpment The Niagara Escarpment is a long escarpment, or cuesta, in Canada and the United States that runs predominantly east–west from New York through Ontario, Michigan, Wisconsin, and into Illinois. The escarpment is most famous as the cliff over ...
geological formations in Central New York (present in 2 counties), southern Ontario (present in 7 counties), and the eastern Upper Peninsula in
Michigan Michigan () is a state in the Great Lakes region of the upper Midwestern United States. With a population of nearly 10.12 million and an area of nearly , Michigan is the 10th-largest state by population, the 11th-largest by area, and the ...
(present in 2 counties). Exceptions are
disjunct population In biology, a taxon with a disjunct distribution is one that has two or more groups that are related but considerably separated from each other geographically. The causes are varied and might demonstrate either the expansion or contraction of a s ...
s that exist in
Alabama (We dare defend our rights) , anthem = "Alabama (state song), Alabama" , image_map = Alabama in United States.svg , seat = Montgomery, Alabama, Montgomery , LargestCity = Huntsville, Alabama, Huntsville , LargestCounty = Baldwin County, Al ...
(in
Fern Cave National Wildlife Refuge Fern Cave National Wildlife Refuge is a National Wildlife Refuge located in northeastern Alabama, near Paint Rock, Alabama in Jackson County. Despite receiving more than 1,200 visitors per year, the facility is unstaffed; it is administered b ...
, a wildlife refuge centered around an off-limits cave in
Jackson County, Alabama Jackson County is the northeasternmost county in the U.S. state of Alabama. As of the 2020 census, the population was 52,579. The county seat is Scottsboro. The county was named for Andrew Jackson, general in the United States Army and afterw ...
, 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 Tennessee ( , ), officially the State of Tennessee, is a landlocked state in the Southeastern region of the United States. Tennessee is the 36th-largest by area and the 15th-most populous of the 50 states. It is bordered by Kentucky to th ...
(in just a single county); these southern populations are at dire risk of
extirpation Local extinction, also known as extirpation, refers to a species (or other taxon) of plant or animal that ceases to exist in a chosen geographic area of study, though it still exists elsewhere. Local extinctions are contrasted with global extinct ...
. An introduced population descended from New York plants is found in
New Jersey New Jersey is a state in the Mid-Atlantic and Northeastern regions of the United States. It is bordered on the north and east by the state of New York; on the east, southeast, and south by the Atlantic Ocean; on the west by the Delaware ...
; it is a remnant of a 1936 effort to practice ex-situ conservation of populations in New York. *In 2020, a new population of hart's-tongue ferns was discovered inside of 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.


Habitat

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 The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
climates.


Conservation


United States

In the United States, ''A. scolopendrium'' var. ''americanum'' was declared endangered by the U.S. Fish and Wildlife Service in 1989. The reasons for its rarity are currently being researched, with
reintroduction Species reintroduction is the deliberate release of a species into the wild, from captivity or other areas where the organism is capable of survival. The goal of species reintroduction is to establish a healthy, genetically diverse, self-sustainin ...
programs in New York and elsewhere also in development.


Canada

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.


Europe

In spite of being much more common in Europe than in North America (and therefore present in more protected areas), ''A. scolopendrium'' is still declining in certain areas of the continent. The fern was listed as "Vulnerable" in the National Red Lists for
Albania Albania ( ; sq, Shqipëri or ), or , also or . officially the Republic of Albania ( sq, Republika e Shqipërisë), is a country in Southeastern Europe. It is located on the Adriatic and Ionian Seas within the Mediterranean Sea and shares ...
in 2014 and
Norway Norway, officially the Kingdom of Norway, is a Nordic country in Northern Europe, the mainland territory of which comprises the western and northernmost portion of the Scandinavian Peninsula. The remote Arctic island of Jan Mayen and t ...
in 2010 (under Criterion D1); considered "critically threatened and rare" in the
Czech Republic The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The ...
's 2012 plant Red List; and "Endangered" in
Sweden Sweden, formally the Kingdom of Sweden,The United Nations Group of Experts on Geographical Names states that the country's formal name is the Kingdom of SwedenUNGEGN World Geographical Names, Sweden./ref> is a Nordic country located on ...
's 2010 Red List. However, it was not considered threatened in
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
's 1996 Red List of Threatened Plants. ''A. scolopendrium'' is protected by law in the
Netherlands ) , anthem = ( en, "William of Nassau") , image_map = , map_caption = , subdivision_type = Sovereign state , subdivision_name = Kingdom of the Netherlands , established_title = Before independence , established_date = Spanish Netherl ...
since 1998.


Uses


Cultivation

''Asplenium scolopendrium'' is often grown as an
ornamental plant Ornamental plants or garden plants are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars that i ...
, with several
cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture, ...
s selected with varying frond form, including with frilled frond margins, forked fronds and cristate forms. The species has gained the
Royal Horticultural Society The Royal Horticultural Society (RHS), founded in 1804 as the Horticultural Society of London, is the UK's leading gardening charity. The RHS promotes horticulture through its five gardens at Wisley (Surrey), Hyde Hall (Essex), Harlow Carr (Nort ...
's
Award of Garden Merit The Award of Garden Merit (AGM) is a long-established annual award for plants by the British Royal Horticultural Society (RHS). It is based on assessment of the plants' performance under UK growing conditions. History The Award of Garden Merit ...
, as has the cultivar 'Angustatum'. The American variation 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 variation.


Herbal medicine

This fern was used in the 1800s as a medicinal plant in folk medicine as a
spleen The spleen is an organ found in almost all vertebrates. Similar in structure to a large lymph node, it acts primarily as a blood filter. The word spleen comes .
tonic (hence an archaic name for the genus, "spleenworts") and for other uses.


Gallery

Phyllitis scolopendrium habitat.jpg, Habitat Phyllitis scolopendrium3.jpg, Spiers harts tongue fern.JPG, Asplenium scolopendrium americanum.jpg,


References

* *


Further reading

* Hyde, H. A., Wade, A. E., & Harrison, S. G. (1978). ''Welsh Ferns''. National Museum of Wales. . * 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.


External links


Commentary and video on Hart's Tongue ferns at Lynn Glen, Dalry, Scotland
{{Authority control scolopendrium Garden plants of North America Medicinal plants of North America Plants described in 1753 Ferns of Europe Ferns of Asia Ferns of the Americas Taxa named by Carl Linnaeus