Marattiaceae | |
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Angiopteris evecta frond | |
Scientific classification![]() | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Division: | Polypodiophyta |
Class: | Polypodiopsida |
Subclass: | Marattiidae Klinge[1] |
Order: | Marattiales Link[1] |
Family: | Marattiaceae Kaulf.[1] |
Genera | |
Synonyms | |
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Marattiaceae is the only family of extant (living)ferns in the orderMarattiales.[1][2] In thePteridophyte Phylogeny Group classification of 2016 (PPG I), Marattiales is the only order in the subclassMarattiidae. The family has six genera and about 110 species.[1] Many are different in appearance from other ferns, having large fronds and fleshy rootstocks.
The Marattiaceae diverged from other ferns very early in their evolutionary history and are quite different from many plants familiar to people intemperate zones. Many of them have massive, fleshy rootstocks and the largest knownfronds of any fern. The Marattiaceae is one of two groups of ferns traditionally known aseusporangiate ferns, meaning that thesporangium is formed from a group of cells as opposed to a leptosporangium in which there is a single initial cell. At least two genera,Angiopteris andMarattia, have been reported to undergo monoplastidic meiosis rather than polyplastidic meiosis, and are the only known examples withineuphyllophytes to do so.[3][4]
The large fronds characteristic of the group are most readily found in the genusAngiopteris, native toAustralasia,Madagascar andOceania. These fronds may be up to 9 meters long in the speciesAngiopteris teysmanniana ofJava. In theHawaiian Islands,Costa Rica, andJamaica, the speciesAngiopteris evecta is naturalized, having escaped from botanical gardens, and is considered aninvasive species.[5]Angiopteris is unique among ferns in having explosively dispersed spores, which may contribute to its ability to spread.[6]
Marattia in the strict sense is found in the neotropics and Hawaii with six recognized species. The genusEupodium is alsoneotropical, with three species, and was originally described for the distinctive stalked synangia of some species.
Ptisana is apaleotropical genus, formerly thought to be part ofMarattia. These plants are 2-4 times pinnate, with fronds often comparable in size to those found inAngiopteris. Terminal segments usually have a prominent suture where they attach.[7] The sporangia lack the labiate apertures ofMarattia andEupodium, and synangia are deeply cut. The name of the genus derives from the resemblance of the synangia topearl barley. The king fern,Ptisana salicina, fromNew Zealand and theSouth Pacific and known inMāori as "para" now has been placed in this genus. Sometimes called the potato fern, this is a large fern with an edible fleshyrhizome that is used as a food source by some indigenous peoples.
The East-Asian genusChristensenia, named after the Danish pteridologistCarl Christensen, is an uncommon fern with distinctive fronds resembling ahorse chestnut leaf, hence the speciesChristensenia aesculifolia, meaning horse-chestnut-leavedChristensenia. Despite the relatively diminutive size of plants in this genus, thestomata ofChristensenia are the largest known in the plant kingdom.[8]
The genusDanaea is endemic to theNeotropics. They have bipinnate leaves with opposite pinnae, which are dimorphic, the fertile leaves much contracted, and covered below with sunken, linearsynangia dehiscing via pores.[9]
in thePteridophyte 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 classPolypodiopsida (ferns), to which it is related as shown in thiscladogram, being asister group toPolypodiidae.[1]
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.[2] The class was lowered in rank to the subclass Marattiidae in the 2009 classification ofMark W. Chase andJames L. Reveal,[10] and subsequent systems such as Christenhusz et al. (2011).[11][12] The Pteridophyte Phylogeny Group (2016) classification retains this rank. In that system, Marattiidae ismonotypic and has one order, Marattiales, one family, Marattiaceae, six genera, and an estimated 111 species.[1]
There have long been four traditional extant genera (Angiopteris,Christensenia,Danaea andMarattia), but phylogenetic analysis has determined the genusMarattia to be paraphyletic, and the genus has been split into three genera,Marattia in the strict sense,Eupodium, andPtisana.[7][11] Christenhusz and Chase placedDanaea in subfamily Danaeoideae and the remaining genera in subfamily Marattioideae,[12] but this subfamilial classification was not taken up by PPG I.[1]
This fern group has a long fossil history with many extinct taxa (Psaronius,Asterotheca,Scolecopteris,Eoangiopteris,Qasimia,Marantoidea,Danaeites,Marattiopsis,Ptychocarpus, etc.).
Exploring the phylogeny of the marattialean ferns[7][13] | Fern Tree of Life[14][15] | ||||||||||||||||||
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Six genera are accepted in the PPG I classification:[1]
Several other genera have been named in the Marattiaceae, namely:Archangiopteris,Clementea,Macroglossum,Protangiopteris,Protomarattia andPsilodochea. These are currently treated assynonyms ofAngiopteris.[1]
Marattiaceae are considered one of the most primitive living lineages of ferns. The earliest members of the family appeared during theCarboniferous, over 300 million years ago. The group has an extensive fossil record extending from the Carboniferous into theJurassic, but post-Jurassic records are scarce.[16]