Angiopteris evecta, commonly known as the king fern, giant fern, elephant fern, oriental vessel fern, Madagascar tree fern, or mule's Foot fern, is a very large rainforest fern in the family Marattiaceae native to most parts of Southeast Asia and Oceania. It has a history dating back about 300 million years, and is believed to have the longest fronds of any fern in the world.
Angiopteris evecta is a self-supporting evergreen perennial fern with very large bipinnate fronds. The trunk-like rhizome is massive, measuring up to 1m (3ft 3in) in diameter. The older portions of the rhizome lie on the ground while the newer growth may rise vertically up to 1.2m (3ft 11in) high.
The arching, glossy green fronds, which emerge from the tip of the rhizome, may reach up to 9m (30ft) long and 2.5m (8ft 2in) wide, with the fleshy green petiole (leaf stem) making up 2m (6ft 7in) of that length.[4][5][6][7] They are said to be the longest fern fronds in the world,[8][9][10] and despite their enormous size they have no woody strengthening tissues in the fronds to keep them erect—instead they are supported entirely by the hydraulic pressure of the sap.[11] On either side of the petiole where it arises from the rhizome there are flat, rounded, leathery, ear-shaped stipules, known as "auricles",[12][13] which can measure up to 15cm (5.9in) in diameter.
The fronds are bipinnate with about 9 to 12 pairs of pinnae measuring up to 150cm (59in) long and 45cm (18in) wide. Each pinnae carries about 30 to 40 pairs of pinnules that measure around 13 by 2cm (5.1 by 0.8in), and both the main rachis and the secondary rachillae (midribs) are pulvinate (swollen at the base).[5][14] Sporangia are borne on the underside of the pinnules, very close to the margin, in clusters of 5 to 8 opposite pairs.[6][11] Overall dimensions of this fern can be up to 7m (23ft) high by 16m (52ft) wide.[4][6][7][15][16]
Evolution
Fossilised fronds bearing a distinct similarity to this plant have been found in Paleozoic rocks from every continent,[11] indicating that the species is very primitive and was widespread around 300 million years ago, a time when ferns and their relatives were the dominant plants on the planet.[8][9][10][11] The geographically isolated communities seen today point to favourable climatic conditions being more widespread in the past.[4]
Taxonomy
Angiopteris evecta was originally named and described as Polypodium evectum by Georg Forster in 1786, in his work Florulae Insularum Australium Prodromus.[17] It was moved to the genus Angiopteris in 1794 by Georg Franz Hoffmann, publishing in the journal Commentationes Societatis Regiae Scientiarum Gottingensis.[18][19]A. evecta is the type species of the genus Angiopteris.[20]
Etymology
The genus name comes from the Ancient Greekaggeion, a vessel, and pteris, a fern, and is a reference to the sporangia.[14] The species epithet is the Latin adjective evectus meaning to carry out, bring forth, raise, or elevate.[21]
Synonyms
As of August2021[update], Plants of the World Online lists 73 synonyms for Angiopteris evecta,[22] which are considered by some authorities to potentially be distinct species, calling for a more thorough taxonomic investigation.[5][15][23]
Angiopteris acrocarpa de Vriese
Angiopteris affinis de Vriese
Angiopteris alata Nadeaud
Angiopteris albidopunctulata Rosenst.
Angiopteris amboinensis de Vriese
Angiopteris angustata Miq.
Angiopteris angustifolia C.Presl
Angiopteris ankolana de Vriese
Angiopteris aphanosorus de Vriese
Angiopteris approximata de Vriese
Angiopteris arborescens (Blanco) Merr.
Angiopteris assamica de Vriese
Angiopteris athroocarpa Alderw.
Angiopteris aurata de Vriese
Angiopteris badioneura de Vriese
Angiopteris beecheyana de Vriese
Angiopteris brongniartiana de Vriese
Angiopteris canaliculata Holttum
Angiopteris caudata de Vriese
Angiopteris commutata C.Presl
Angiopteris crassifolia de Vriese
Angiopteris cumingii Hieron.
Angiopteris cupreata de Vriese
Angiopteris cuspidata de Vriese
Angiopteris dregeana de Vriese
Angiopteris durvilleana de Vriese
Angiopteris elongata Hieron.
Angiopteris erecta Desv.
Angiopteris evanidostriata Hieron.
Angiopteris evecta var. rurutensis E.D.Br.
Angiopteris grisea Alderw.
Angiopteris hellwigii Hieron.
Angiopteris inconstans Alderw.
Angiopteris indica Desv.
Angiopteris intricata C.Presl
Angiopteris javanica C.Presl
Angiopteris lancifoliolata Alderw.
Angiopteris lasegueana de Vriese
Angiopteris lauterbachii Hieron.
Angiopteris leytensis Alderw.
Angiopteris longifolia Grev. & Hook.
Angiopteris lorentzii Rosenst.
Angiopteris medogensis Ching & Y.X.Lin
Angiopteris mekongensis Ching ex C.Chr. & Tardieu
Angiopteris microsporangia de Vriese
Angiopteris microura Copel.
Angiopteris miqueliana de Vriese
Angiopteris monstruosa Alderw.
Angiopteris naumannii Hieron.
Angiopteris norrisii Rosenst.
Angiopteris novocaledonica Hieron.
Angiopteris oligotheca Hieron.
Angiopteris olivacea Alderw.
Angiopteris palauensis Hieron.
Angiopteris pallescens de Vriese
Angiopteris pallida Rosenst.
Angiopteris palmiformis (Cav.) C.Chr.
Angiopteris papandayanensis Hieron.
Angiopteris polytheca C.Chr. & Tardieu
Angiopteris presliana de Vriese
Angiopteris ruttenii Alderw.
Angiopteris similis C.Presl
Angiopteris stellatosora C.Chr.
Angiopteris subfurfuracea Alderw.
Angiopteris teysmanniana de Vriese
Angiopteris uncinata de Vriese
Angiopteris willinkii Miquel
Callipteris heterophylla Moore
Clementea palmiformis Cav.
Danaea evecta (G.Forst.) Spreng.
Lomaria pedunculata Goldm.
Myriotheca arborescens Blanco
Polypodium evectum G.Forst.
Distribution and habitat
Angiopteris evecta is native to southeast Asia and Oceania, from Sri Lanka and Bangladesh in the west through to Melanesia, Micronesia and Polynesia in the east, and from Japan in the north to northern and eastern Australia in the south. It has been introduced to most of the rest of tropical Asia, as well as Madagascar and parts of the tropical Americas. It has become naturalised in Hawaii, Jamaica, Costa Rica and Cuba.[4][5][15][22][24][25]
The species grows in rainforest on very rich soils, often of volcanic origin[4][15] and prefers a very warm wet climate. It is usually an understorey plant in well developed rainforest, especially along creek banks in deep sheltered gullies where there is good drainage and a plentiful supply of fresh water, but is occasionally found in more exposed situations.[4][16][26] The preferred annual mean temperature range is 19–27°C (66–81°F) and annual precipitation between 1,000–5,447mm (39–214in). It grows at elevations from sea level to 1,500m (4,900ft).[5][7][15]
Cultural uses
The starchy rhizomes are eaten after long processing to remove toxins, used to perfume coconut oil, to flavour rice and to produce an intoxicating drink.[4][5][7] The 1889 book The Useful Native Plants of Australia records Indigenous Australians ate the pith of this fern.[27]
Conservation
The conservation status of Angiopteris evecta varies from place to place. For example in Australia's Northern Territory it is listed as vulnerable, with only one small population in north eastern Arnhem Land;[9][28] in New South Wales, where suitable habitat is restricted to a small area in the north east corner of the state and only a single, non-reproductive specimen is known,[4] it is listed as endangered.[16][28] However in the state of Queensland, which lies in between the other two states and where there is an abundance of suitable habitat, it is listed as least concern.[1][16][28]
As of 16August2021[update], this species has not been assessed by the IUCN.
Invasive potential
When introduced to an area with a suitable climate, Angiopteris evecta can establish dense stands that inhibit local species.[29] It is listed as invasive in Costa Rica, Cuba, Jamaica and Hawaii, where in each case it has escaped from plantings in botanic gardens. It has also been introduced to many tropical countries and has repeatedly escaped from cultivation.[5]
Gallery
Pulvini on the rachillae (midribs of lateral branches)
Growing in a steep gully on the Lamb Range, Queensland, Australia
On the Alexandra Range near Cape Tribulation, Australia
Underside of a frond
Growing at Garfield Park Conservatory. The swollen bases of the petioles are clearly seen here, as are the rounded stipules
Underside of pinnules, showing sporangia and the swollen basal attachment. Royal Botanic Garden, Sydney
In Cairns Botanic Garden, August 2021. This image illustrates the prostrate older section of the rhizome and the erect younger portion
"Angiopteris evecta - International Plant Names Index". International Plant Names Index. The Royal Botanic Gardens, Kew, Harvard University Herbaria & Libraries and Australian National Botanic Gardens. Retrieved 5 December 2020.
"Short Cuts". Australian Native Plants Society (Australia). Australian Native Plants Society (Australia). Retrieved 6 December 2020.
Camus, Josephine M. (2020). "Angiopteris". Flora of Australia. Australian Biological Resources Study, Department of Agriculture, Water and the Environment: Canberra. Retrieved 18 August 2021.
"Polypodium evectum". Australian Plant Name Index (APNI). Centre for Australian National Biodiversity Research, Australian Government. Retrieved 16 October 2020.
"Angiopteris evecta". Australian Plant Name Index (APNI). Centre for Australian National Biodiversity Research, Australian Government. Retrieved 16 October 2020.
"Angiopteris - International Plant Names Index". International Plant Names Index. The Royal Botanic Gardens, Kew, Harvard University Herbaria & Libraries and Australian National Botanic Gardens. Retrieved 5 December 2020.
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