Marattiaceae Temporal range: | |
---|---|
![]() | |
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 | |
|
Marattiaceae is the only family of extant (living) ferns in the order Marattiales. [1] [2] 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. [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 in temperate zones. 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 is formed from a group of cells as opposed to a leptosporangium in which there is a single initial cell.
It is the only known extant group of euphyllophytes which undergoes monoplastidic meiosis rather than polyplastidic meiosis, and is considered a primitive trait. [3]
The large fronds characteristic of the group are most readily found in the genus Angiopteris , native to Australasia, Madagascar and Oceania. These fronds may be up to 9 meters long in the species Angiopteris teysmanniana of Java. In Jamaica the species Angiopteris evecta is widely naturalized and is registered as an invasive species. The plant was introduced by Captain Bligh from Tahiti as a staple food for slaves and cultivated in the Castleton Botanical Garden in 1860. From there it was able to distribute itself throughout the eastern half of the island.
Marattia in the strict sense is found in the neotropics and Hawaii. The genus Eupodium is also neotropical, 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 is a paleotropical genus. These plants are 2-4 times pinnate, with fronds often comparable in size to those found in Angiopteris. 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. The king fern, Ptisana salicina , from New Zealand and the South Pacific and known in Māori as "para" now has been placed in this genus. Sometimes called the potato fern, this is a large fern with an edible fleshy rhizome 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 of Christensenia are the largest known in the plant kingdom. [4]
The genus Danaea is endemic to the Neotropics. 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. [5]
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, being a sister group to Polypodiidae. [1]
Polypodiopsida |
| ||||||||||||||||||
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 of Mark W. Chase and James L. Reveal, [6] and subsequent systems such as Christenhusz et al. (2011). [7] [8] The Pteridophyte Phylogeny Group (2016) classification retains this rank. In that system, Marattiidae is monotypic 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 and Marattia ), 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 , and Ptisana . [9] [7] Christenhusz and Chase placed Danaea in subfamily Danaeoideae and the remaining genera in subfamily Marattioideae, [8] 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 [9] [10] | Fern Tree of Life [11] [12] | ||||||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
|
|
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 and Psilodochea. These are currently treated as synonyms of Angiopteris. [1]
Marattiaceae are considered one of the most primitive living lineages of ferns. The earliest members of the family appeared during the Carboniferous, over 300 million years ago. The group has an extensive fossil record extending from the Carboniferous into the Jurassic, but post-Jurassic records are scarce. [13]
The ferns are a group of vascular plants that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissues that conduct water and nutrients and in having life cycles in which the branched sporophyte is the dominant phase.
Ophioglossaceae, the adder's-tongue family, is a small family of ferns. In the Pteridophyte Phylogeny Group classification of 2016, it is the only family in the order Ophioglossales, which together with the Psilotales is placed in the subclass Ophioglossidae. The Ophioglossidae are one of the groups traditionally known as eusporangiate ferns. Members of the family differ from other ferns in a number of ways. Many have only a single fleshy leaf at a time. Their gametophytes are subterranean and rely on fungi for energy.
Osmunda is a genus of primarily temperate-zone ferns of family Osmundaceae. Five to ten species have been listed for this genus.
The order Polypodiales encompasses the major lineages of polypod ferns, which comprise more than 80% of today's fern species. They are found in many parts of the world including tropical, semitropical and temperate areas.
A pteridophyte is a vascular plant that reproduces by means of spores. Because pteridophytes produce neither flowers nor seeds, they are sometimes referred to as "cryptogams", meaning that their means of reproduction is hidden.
Psilotaceae is a family of ferns consisting of two genera, Psilotum and Tmesipteris with about a dozen species. It is the only family in the order Psilotales.
Oleandra is a genus of ferns. In the Pteridophyte Phylogeny Group classification of 2016, it is the only genus in the family Oleandraceae, which is placed in suborder Polypodiineae, order Polypodiales. Alternatively, the family may be placed in a very broadly defined family Polypodiaceae sensu lato as the subfamily Oleandroideae.
Nephrolepis is a genus of about 30 species of ferns. It is the only genus in the family Nephrolepidaceae, placed in the suborder Aspleniineae of the order Polypodiales in the Pteridophyte Phylogeny Group classification of 2016. The genus is commonly referred to as macho ferns or Boston ferns. The fronds are long and narrow, and once-pinnate, in the case of one Bornean species reaching thirty feet in length.
The Athyriaceae are a family of terrestrial ferns in the order Polypodiales. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family is placed in the suborder Aspleniineae, and includes two genera. Alternatively, it may be treated as the subfamily Athyrioideae of a very broadly defined family Aspleniaceae. The family has with a cosmopolitan distribution.
Eusporangiate ferns are vascular spore plants, whose sporangia arise from several epidermal cells and not from a single cell as in leptosporangiate ferns. Typically these ferns have reduced root systems and sporangia that produce large amounts of spores.
The Polypodiidae, commonly called leptosporangiate ferns, formerly Leptosporangiatae, are one of four subclasses of ferns, the largest of these being the largest group of living ferns, including some 11,000 species worldwide. The group has also been treated as the class Pteridopsida or Polypodiopsida, although other classifications assign them a different rank. Older names for the group include Filicidae and Filicales, although at least the "water ferns" were then treated separately.
Saccoloma is a fern genus in family Saccolomataceae. It is the only genus in the family in the Pteridophyte Phylogeny Group classification of 2016, but further investigation is needed. It is pantropical and its species are found in wet, shaded forest areas. Saccoloma species are characterized by an omega-shaped (Ω) vascular bundle in the cross-sections of their petioles. The common name soralpouch fern is used for Saccoloma.
Saccolomataceae is a family of ferns in the order Polypodiales with about 19 species. It has been formerly treated as part of the Dennstaedtiaceae, however it has been classified as its own family according to Smith et al. (2006) The genus Saccoloma has been classified to include Orthiopteris, but the phylogeny of the group not yet fully understood. The family includes a dozen known species.
Didymochlaena is a genus of ferns. In the Pteridophyte Phylogeny Group classification of 2016, it is the only genus in the family Didymochlaenaceae. Alternatively, the family may be placed in a very broadly defined family Polypodiaceae sensu lato as the subfamily Didymochlaenoideae.
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, which supported recognition of this group on the basis of genetic analysis and morphology. Ptisana can be distinguished from Marattia by the presence of distinct sutures at the point of leaflet attachment, deeply cut synangia, and the absence of labiate sporangial apertures. The name Ptisana is derived from the Latin word for pearl barley, an allusion to the shape of the synangia.
Eupodium is a genus of ferns in the family Marattiaceae native to the Neotropics. Traditionally, many taxonomists have included Eupodium within the genus Marattia. However, molecular phylogenetic studies and morphological studies of extant and fossil taxa support the recognition of Eupodium as a lineage distinct from Marattia. Morphologically, Eupodium was thought to be distinct among the Marattiaceae in only having one frond at a time, bearing awns along veins, and having stalked synangia. However, recent phylogenetic work found that an additional species native to Brazil, Eupodium cicutifolium, which lacks these characters, is also genetically in Eupodium, making the genus challenging to distinguish morphologically. Eupodium cicutifolium does occasionally have short-stalked synangia, and has spinulose spores like the other Eupodium species, but is otherwise easily mistaken for a Marattia or Ptisana. The genus currently has four recognized species.
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 that Marattia in the broad sense was paraphyletic, and subsequently the genera Ptisana and Eupodium were split off. Except for one species in Hawaii, the genus is neotropical.
Tectariaceae is a family of leptosporangiate ferns in the order Polypodiales. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family is placed in the suborder Polypodiineae. Alternatively, it may be treated as the subfamily Tectarioideae of a very broadly defined family Polypodiaceae sensu lato. The family comprises seven genera, of which Tectaria is by far the largest.
Metaxya is a neotropical genus of ferns in the order Cyatheales. It is the only genus in the family Metaxyaceae in the Pteridophyte Phylogeny Group classification of 2016. Alternatively, the genus may be placed in the subfamily Metaxyoideae of a more broadly defined family Cyatheaceae, the family placement used in Plants of the World Online as of November 2019.
Ophioglossidae is one of the four subclasses of Polypodiopsida (ferns). This subclass consists of the ferns commonly known as whisk ferns, grape ferns, adder's-tongues and moonworts. It is equivalent to the class Psilotopsida in previous treatments, including Smith et al. (2006). The subclass contains two orders, Psilotales and Ophioglossales, whose relationship was only confirmed by molecular phylogenetic studies.