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''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 Rainforests are characterized by a closed and continuous tree canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforest can be classified as tropical rainforest or temperate rainfores ...
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 family
Marattiaceae Marattiaceae is the only family of extant (living) ferns in the order Marattiales. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), Marattiales is the only order in the subclass Marattiidae. The family has six genera and about ...
native to most parts of
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical United Nations geoscheme for Asia#South-eastern Asia, south-eastern region of Asia, consistin ...
and
Oceania Oceania (, , ) is a region, geographical region that includes Australasia, Melanesia, Micronesia, and Polynesia. Spanning the Eastern Hemisphere, Eastern and Western Hemisphere, Western hemispheres, Oceania is estimated to have a land area of ...
. It has a history dating back about 300 million years, and is believed to have the longest fronds of any fern in the world.


Description

''Angiopteris evecta'' is a self-supporting
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 ...
perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
fern with very large
bipinnate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular o ...
fronds. The trunk-like
rhizome In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow hori ...
is massive, measuring up to in diameter. The older portions of the rhizome lie on the ground while the newer growth may rise vertically up to high. The arching, glossy green
frond A frond is a large, divided leaf. In both common usage and botanical nomenclature, the leaves of ferns are referred to as fronds and some botanists restrict the term to this group. Other botanists allow the term frond to also apply to the lar ...
s, which emerge from the tip of the rhizome, may reach up to long and wide, with the fleshy green petiole (leaf stem) making up of that length. They are said to be the longest fern fronds in the world, 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 Hydraulics (from Greek: Υδραυλική) is a technology and applied science using engineering, chemistry, and other sciences involving the mechanical properties and use of liquids. At a very basic level, hydraulics is the liquid coun ...
of the sap. On either side of the petiole where it arises from the rhizome there are flat, rounded, leathery, ear-shaped
stipule In botany, a stipule is an outgrowth typically borne on both sides (sometimes on just one side) of the base of a leafstalk (the petiole). Stipules are considered part of the anatomy of the leaf of a typical flowering plant, although in many speci ...
s, known as "auricles", which can measure up to in diameter. The fronds are
bipinnate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular o ...
with about 9 to 12 pairs of pinnae measuring up to long and wide. Each pinnae carries about 30 to 40 pairs of pinnules that measure around , and both the main rachis and the secondary rachillae (midribs) are pulvinate (swollen at the base). Sporangia are borne on the underside of the pinnules, very close to the margin, in clusters of 5 to 8 opposite pairs. Overall dimensions of this fern can be up to high by wide.


Evolution

Fossilised fronds bearing a distinct similarity to this plant have been found in
Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ' ...
rocks from every continent, 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. The geographically isolated communities seen today point to favourable climatic conditions being more widespread in the past.


Taxonomy

''Angiopteris evecta'' was originally named and described as ''Polypodium evectum'' by
Georg Forster Johann George Adam Forster, also known as Georg Forster (, 27 November 1754 – 10 January 1794), was a German naturalist, ethnologist, travel writer, journalist and revolutionary. At an early age, he accompanied his father, Johann Reinhold F ...
in 1786, in his work '' Florulae Insularum Australium Prodromus''. It was moved to the genus Angiopteris in 1794 by
Georg Franz Hoffmann Georg Franz Hoffmann was a German botanist and lichenologist. He was born on 13 January 1760 in Marktbreit, Germany, and died on 17 March 1826 in Moscow, Russia. Professional career After graduating from the University of Erlangen in 1786, he w ...
, publishing in the journal '' Commentationes Societatis Regiae Scientiarum Gottingensis''. ''A. evecta'' is the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
of the genus ''
Angiopteris ''Angiopteris'' is a genus of huge evergreen ferns from the family Marattiaceae, found throughout the paleotropics from Madagascar to the South Pacific islands. Species of smaller stature with elongate synangia and creeping rhizomes are sometime ...
''.


Etymology

The genus name comes from the
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
''aggeion'', a vessel, and ''pteris'', a fern, and is a reference to the sporangia. The species epithet is the Latin adjective ''evectus'' meaning to carry out, bring forth, raise, or elevate.


Synonyms

,
Plants of the World Online Plants of the World Online (POWO) is an online database published by the Royal Botanic Gardens, Kew. It was launched in March 2017 with the ultimate aim being "to enable users to access information on all the world's known seed-bearing plants by ...
lists 73 synonyms for ''Angiopteris evecta'', which are considered by some authorities to potentially be distinct species, calling for a more thorough taxonomic investigation. * ''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 Sri Lanka (, ; si, ශ්‍රී ලංකා, Śrī Laṅkā, translit-std=ISO (); ta, இலங்கை, Ilaṅkai, translit-std=ISO ()), formerly known as Ceylon and officially the Democratic Socialist Republic of Sri Lanka, is an ...
and
Bangladesh Bangladesh (}, ), officially the People's Republic of Bangladesh, is a country in South Asia. It is the eighth-most populous country in the world, with a population exceeding 165 million people in an area of . Bangladesh is among the mos ...
in the west through to
Melanesia Melanesia (, ) is a subregion of Oceania in the southwestern Pacific Ocean. It extends from Indonesia's New Guinea in the west to Fiji in the east, and includes the Arafura Sea. The region includes the four independent countries of Fiji, Va ...
,
Micronesia Micronesia (, ) is a subregion of Oceania, consisting of about 2,000 small islands in the western Pacific Ocean. It has a close shared cultural history with three other island regions: the Philippines to the west, Polynesia to the east, and ...
and
Polynesia Polynesia () "many" and νῆσος () "island"), to, Polinisia; mi, Porinihia; haw, Polenekia; fj, Polinisia; sm, Polenisia; rar, Porinetia; ty, Pōrīnetia; tvl, Polenisia; tkl, Polenihia (, ) is a subregion of Oceania, made up of ...
in the east, and from
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
in the north to northern and eastern
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
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 Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only stat ...
,
Jamaica Jamaica (; ) is an island country situated in the Caribbean Sea. Spanning in area, it is the third-largest island of the Greater Antilles and the Caribbean (after Cuba and Hispaniola). Jamaica lies about south of Cuba, and west of His ...
,
Costa Rica Costa Rica (, ; ; literally "Rich Coast"), officially the Republic of Costa Rica ( es, República de Costa Rica), is a country in the Central American region of North America, bordered by Nicaragua to the north, the Caribbean Sea to the no ...
and
Cuba Cuba ( , ), officially the Republic of Cuba ( es, República de Cuba, links=no ), is an island country comprising the island of Cuba, as well as Isla de la Juventud and several minor archipelagos. Cuba is located where the northern Caribbea ...
. The species grows in
rainforest Rainforests are characterized by a closed and continuous tree canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforest can be classified as tropical rainforest or temperate rainfores ...
on very rich soils, often of volcanic origin and prefers a very warm wet climate. It is usually an
understorey In forestry and ecology, understory (American English), or understorey (Commonwealth English), also known as underbrush or undergrowth, includes plant life growing beneath the forest canopy without penetrating it to any great extent, but abov ...
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. The preferred annual mean temperature range is and annual precipitation between . It grows at elevations from sea level to .


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. The 1889 book ''The Useful Native Plants of Australia'' records
Indigenous Australians Indigenous Australians or Australian First Nations are people with familial heritage from, and membership in, the ethnic groups that lived in Australia before British colonisation. They consist of two distinct groups: the Aboriginal peoples ...
ate the pith of this fern.


Conservation

The conservation status of ''Angiopteris evecta'' varies from place to place. For example in Australia's
Northern Territory The Northern Territory (commonly abbreviated as NT; formally the Northern Territory of Australia) is an states and territories of Australia, Australian territory in the central and central northern regions of Australia. The Northern Territory ...
it is listed as vulnerable, with only one small population in north eastern
Arnhem Land Arnhem Land is a historical region of the Northern Territory of Australia, with the term still in use. It is located in the north-eastern corner of the territory and is around from the territory capital, Darwin. In 1623, Dutch East India Compan ...
; in
New South Wales ) , nickname = , image_map = New South Wales in Australia.svg , map_caption = Location of New South Wales in AustraliaCoordinates: , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , es ...
, 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, it is listed as
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inva ...
. However in the state of
Queensland ) , nickname = Sunshine State , image_map = Queensland in Australia.svg , map_caption = Location of Queensland in Australia , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , established_ ...
, which lies in between the other two states and where there is an abundance of suitable habitat, it is listed as
least concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...
. , this species has not been assessed by the
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
.


Invasive potential

When introduced to an area with a suitable climate, ''Angiopteris evecta'' can establish dense stands that inhibit local species. It is listed as invasive in
Costa Rica Costa Rica (, ; ; literally "Rich Coast"), officially the Republic of Costa Rica ( es, República de Costa Rica), is a country in the Central American region of North America, bordered by Nicaragua to the north, the Caribbean Sea to the no ...
,
Cuba Cuba ( , ), officially the Republic of Cuba ( es, República de Cuba, links=no ), is an island country comprising the island of Cuba, as well as Isla de la Juventud and several minor archipelagos. Cuba is located where the northern Caribbea ...
,
Jamaica Jamaica (; ) is an island country situated in the Caribbean Sea. Spanning in area, it is the third-largest island of the Greater Antilles and the Caribbean (after Cuba and Hispaniola). Jamaica lies about south of Cuba, and west of His ...
and
Hawaii Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only stat ...
, 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.


Gallery

File: Angiopteris evecta 13zz.jpg , Pulvini on the rachillae (midribs of lateral branches) File: Angiopteris-evecta-lamb-range1.jpg , Growing in a steep gully on the
Lamb Range Lamb Range is a locality in the Cairns Region, Queensland, Australia. In the , Lamb Range had a population of 0. Geography The locality takes its name from the mountain range Lamb Range. The locality that lies to the west of most of the south ...
, Queensland, Australia File: Angiopteris-evecta-alexandra-range1.jpg , On the Alexandra Range near
Cape Tribulation Cape Tribulation is a headland and coastal locality in the Shire of Douglas in northern Queensland, Australia. In the , Cape Tribulation had a population of 118 people. Geography The locality is north of Cairns. It is within the Daintree Na ...
, Australia File: Angiopteris evecta kz09.jpg , Underside of a frond File: Angiopteris evecta kz1.jpg , Growing at
Garfield Park Conservatory Garfield Park Conservatory, located in Garfield Park in Chicago, is one of the largest greenhouse conservatories in the United States. Often referred to as "landscape art under glass", the Garfield Park Conservatory occupies approximately insi ...
. The swollen bases of the petioles are clearly seen here, as are the rounded stipules File: Angiopteris evecta, Royal Botanic Garden Sydney IMG 20190502 115445.jpg , Underside of pinnules, showing sporangia and the swollen basal attachment.
Royal Botanic Garden, Sydney The Royal Botanic Garden, Sydney is a heritage-listed major botanical garden, event venue and public recreation area located at Farm Cove on the eastern fringe of the Sydney central business district, in the City of Sydney local government a ...
File: Angiopteris evecta SF21231 01.jpg , In Cairns Botanic Garden, August 2021. This image illustrates the prostrate older section of the rhizome and the erect younger portion


References


External links

* *
View a map
of recorded sightings of ''Angiopteris evecta'' at the
Australasian Virtual Herbarium The ''Australasian Virtual Herbarium'' (AVH) is an online resource that allows access to plant specimen data held by various Australian and New Zealand herbaria. It is part of the Atlas of Living Australia (ALA), and was formed by the amalgamat ...

See images
of ''Angiopteris evecta'' on Flickriver {{Taxonbar, from=Q4763286 Marattiidae Flora of New South Wales Flora of Queensland Flora of the Northern Territory Flora of Papua New Guinea Flora of the Maluku Islands Wet Tropics of Queensland Flora of Polynesia Flora of Singapore Flora of Melanesia Flora of Micronesia Flora of Sumatra Flora of Java Flora of the Lesser Sunda Islands Flora of Sulawesi Flora of Borneo