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Marattiaceae is the only family of extant (living)
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 exce ...
s 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 110 species. Many are different in appearance from other ferns, having large fronds and fleshy rootstocks.


Description

The Marattiaceae diverged from other ferns very early in their evolutionary history and are quite different from many plants familiar to people in
temperate zones In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...
. Many of them have massive, fleshy rootstocks and the largest known fronds of any fern. The Marattiaceae is one of two groups of ferns traditionally known as eusporangiate ferns, meaning that the
sporangium A sporangium (; from Late Latin, ) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their life cy ...
is formed from a group of cells as opposed to a leptosporangium in which there is a single initial cell. The large fronds characteristic of the group are most readily found in 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 some ...
'', native to
Australasia Australasia is a region that comprises Australia, New Zealand and some neighbouring islands in the Pacific Ocean. The term is used in a number of different contexts, including geopolitically, physiogeographically, philologically, and ecologic ...
,
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Afric ...
and
Oceania Oceania (, , ) is a geographical region that includes Australasia, Melanesia, Micronesia, and Polynesia. Spanning the Eastern and Western hemispheres, Oceania is estimated to have a land area of and a population of around 44.5 million ...
. These fronds may be up to 9 meters long in the species '' Angiopteris teysmanniana'' of
Java Java (; id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 151.6 million people, Java is the world's mo ...
. In
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 Hispa ...
the species ''
Angiopteris evecta ''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 A ...
'' is widely naturalized and is registered as an
invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species adv ...
. The plant was introduced by Captain Bligh from
Tahiti Tahiti (; Tahitian ; ; previously also known as Otaheite) is the largest island of the Windward group of the Society Islands in French Polynesia. It is located in the central part of the Pacific Ocean and the nearest major landmass is Austra ...
as a staple food for
slaves Slavery and enslavement are both the state and the condition of being a slave—someone forbidden to quit one's service for an enslaver, and who is treated by the enslaver as property. Slavery typically involves slaves being made to perf ...
and cultivated in the
Castleton Botanical Garden Castleton Botanical Garden is a horticultural site of interest established in 1862 and located 19 miles from Kingston, Jamaica.Postcard, undated. Text "Greetings from Jamaica. Castleton Gardens, 19 miles from Kingston".Postcard, undated. Text "C ...
in 1860. From there it was able to distribute itself throughout the eastern half of the island. ''
Marattia ''Marattia'' is a small genus of primitive, large, fleshy eusporangiate ferns. It is the type genus of the family Marattiaceae, order Marattiales and class Marattiopsida. Formerly considered to be a much larger genus, genetic analysis has shown ...
'' in the strict sense is found in the neotropics and Hawaii. The genus ''
Eupodium ''Eupodium'' is a genus of ferns in the family Marattiaceae native to the Neotropics. Traditionally, many taxonomists have included ''Eupodium'' within the genus ''Marattia'' (along with ''Ptisana''). However, molecular phylogenetic studies and m ...
'' is also
neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In bioge ...
, with three species. It has fronds that are 2-5 times pinnate, distinctive stalked synangia, and awns on distal blade segments. Blade division decreases towards the apex of the frond. Plants of ''Eupodium'' usually only have one frond per plant per year (sometimes two). ''
Ptisana Sori of ''Ptisana fraxinea'' ''Ptisana'' is a genus in the eusporangiate fern family Marattiaceae, comprising species historically treated in the genus ''Marattia''. The establishment of this genus follows the 2008 work by Andrew G. Murdock, whi ...
'' is a
paleotropical The Paleotropical Kingdom (Paleotropis) is a floristic kingdom comprising tropical areas of Africa, Asia and Oceania (excluding Australia and New Zealand), as proposed by Ronald Good and Armen Takhtajan. Part of its flora, inherited from the ancien ...
genus. These plants are 2-4 times pinnate, with fronds often comparable in size to those found in
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 some ...
. Terminal segments usually have a prominent suture where they attach. The sporangia lack the labiate apertures of ''Marattia'' and ''Eupodium'', and synangia are deeply cut. The name of the genus derives from the resemblance of the synangia to
pearl barley Pearl barley, or pearled barley, is barley that has been processed to remove its fibrous outer hull and polished to remove some or all of the bran layer. It is the most common form of barley for human consumption because it cooks faster and i ...
. The king fern, ''
Ptisana salicina ''Ptisana salicina'', or king fern, is a species of fern native to Norfolk Island, New Zealand and the South Pacific. Large and robust with a distinctive tropical appearance, it has fronds up to 5 metres (16 feet +/-) tall that arise from a st ...
'', from
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island coun ...
and the
South Pacific The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the continen ...
and known in
Māori Māori or Maori can refer to: Relating to the Māori people * Māori people of New Zealand, or members of that group * Māori language, the language of the Māori people of New Zealand * Māori culture * Cook Islanders, the Māori people of the Co ...
as "para" now has been placed in this genus. Sometimes called the potato fern, this is a large fern with an edible fleshy
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 ...
that is used as a food source by some indigenous peoples. The East-Asian genus '' Christensenia'' is named in honor of the Danish pteridologist Carl Christensen is an uncommon fern with distinctive fronds resembling a horse chestnut leaf, hence the species '' Christensenia aesculifolia'', meaning horse-chestnut-leaved ''Christensenia''. Despite the relatively diminutive size of plants in this genus, the
stomata In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is b ...
of ''Christensenia'' are the largest known in the plant kingdom. The genus '' Danaea'' is endemic to the
Neotropics The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In bioge ...
. They have bipinnate leaves with opposite pinnae, which are dimorphic, the fertile leaves much contracted, and covered below with sunken, linear synangia dehiscing via pores.


Taxonomy

in the Pteridophyte Phylogeny Group classification of 2016 (PPG I), Marattiaceae is the only family in the order Marattiales, which in turn is the only order in the subclass Marattiidae. Marattiidae is one of four subclasses of class Polypodiopsida (ferns), to which it is related as shown in this
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
, being a
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to Polypodiidae.


History of classification

In the molecular phylogenetic classification of Smith et al. in 2006, the Marattiales formed the single member of the class Marattiopsida. Four genera were recognized. The class was lowered in rank to the subclass Marattiidae in the 2009 classification of
Mark W. Chase Mark Wayne Chase (born 1951) is a US-born British botanist. He is noted for work in plant classification and evolution, and one of the instigators of the Angiosperm Phylogeny Group-classification for flowering plants which is partly based on DNA ...
and James L. Reveal, and subsequent systems such as Christenhusz et al. (2011). The Pteridophyte Phylogeny Group (2016) classification retains this rank. In that system, Marattiidae is
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispe ...
and has one order, Marattiales, one family, Marattiaceae, six genera, and an estimated 111 species. There have long been four traditional extant genera (''
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 some ...
'', '' Christensenia'', '' Danaea'' and ''
Marattia ''Marattia'' is a small genus of primitive, large, fleshy eusporangiate ferns. It is the type genus of the family Marattiaceae, order Marattiales and class Marattiopsida. Formerly considered to be a much larger genus, genetic analysis has shown ...
''), but phylogenetic analysis has determined the genus ''Marattia'' to be paraphyletic, and the genus has been split into three genera, ''Marattia'' in the strict sense, ''
Eupodium ''Eupodium'' is a genus of ferns in the family Marattiaceae native to the Neotropics. Traditionally, many taxonomists have included ''Eupodium'' within the genus ''Marattia'' (along with ''Ptisana''). However, molecular phylogenetic studies and m ...
'', and ''
Ptisana Sori of ''Ptisana fraxinea'' ''Ptisana'' is a genus in the eusporangiate fern family Marattiaceae, comprising species historically treated in the genus ''Marattia''. The establishment of this genus follows the 2008 work by Andrew G. Murdock, whi ...
''. Christenhusz and Chase placed ''Danaea'' in subfamily Danaeoideae and the remaining genera in subfamily Marattioideae, but this subfamilial classification was not taken up by PPG I. This fern group has a long fossil history with many extinct taxa ('' Psaronius'', '' Asterotheca'', '' Scolecopteris'', '' Eoangiopteris'', '' Qasimia'', '' Marantoidea'', '' Danaeites'', '' Marattiopsis'', etc.).


Genera

Six genera are accepted in the PPG I classification: *''
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 some ...
'' Hoffm. *'' Christensenia'' Maxon *'' Danaea'' Sm. *''
Eupodium ''Eupodium'' is a genus of ferns in the family Marattiaceae native to the Neotropics. Traditionally, many taxonomists have included ''Eupodium'' within the genus ''Marattia'' (along with ''Ptisana''). However, molecular phylogenetic studies and m ...
'' J.Sm. *''
Marattia ''Marattia'' is a small genus of primitive, large, fleshy eusporangiate ferns. It is the type genus of the family Marattiaceae, order Marattiales and class Marattiopsida. Formerly considered to be a much larger genus, genetic analysis has shown ...
'' Sw. *''
Ptisana Sori of ''Ptisana fraxinea'' ''Ptisana'' is a genus in the eusporangiate fern family Marattiaceae, comprising species historically treated in the genus ''Marattia''. The establishment of this genus follows the 2008 work by Andrew G. Murdock, whi ...
'' Murdock Several other genera have been named in the Marattiaceae, namely: ''Archangiopteris'', ''Clementea'', ''Macroglossum'', ''Protangiopteris'', ''Protomarattia'' and ''Psilodochea''. These are currently treated as
synonyms A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are ...
of ''Angiopteris''.


Evolutionary history

Marattiaceae are considered one of the most primitive living lineages of ferns. The earliest members of the family appeared during the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferou ...
, over 300 million years ago. The group has an extensive fossil record extending from the Carboniferous into the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
, but post-Jurassic records are scarce.


References


External links


Tree of Life Marattiaceae
{{Taxonbar, from1=Q178286, from2=Q15708883, from3=Q6755062, from4=Q12967693 Marattiidae Fern families