Leptosporangiate fern | |
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Pteridium aquilinum | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Division: | Polypodiophyta |
Class: | Polypodiopsida |
Subclass: | Polypodiidae Cronquist, Takht. & W.Zimm. [1] |
Orders | |
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. [2] [3] [4] The group has also been treated as the class Pteridopsida or Polypodiopsida, [5] although other classifications assign them a different rank. [6] Older names for the group include Filicidae and Filicales, although at least the "water ferns" (now the Salviniales) were then treated separately.
The leptosporangiate ferns are one of the four major groups of ferns, with the other three being the eusporangiate ferns comprising the marattioid ferns (Marattiidae, Marattiaceae), the horsetails (Equisetiidae, Equisetaceae), and whisk ferns and moonworts. [4] [5] There are approximately 8465 species of living leptosporangiate ferns, compared with about 2070 for all other ferns, totalling 10535 species of ferns. [3] Almost a third of leptosporangiate fern species are epiphytes. [7]
These ferns are called leptosporangiate because their sporangia arise from a single epidermal cell and not from a group of cells as in eusporangiate ferns (a polyphyletic lineage). The mature sporangia have a wall that is just a single cell thick, [8] and are typically covered with a scale called the indusium, which can cover the whole sorus, forming a ring or cup around the sorus, or can also be strongly reduced to completely absent. Many leptosporangiate ferns have an annulus around the sporangium, which ejects the spores. [9]
The leptosporangiate ferns were first recognized as a group, the "Leptosporangiateen", by Karl Ritter von Goebel in 1881, who placed the eusporangiate ferns with seed plants and vascular plants into a coeval "Eusporangiateen". As this classification artificially split the ferns, Christian Luerssen subdivided the homosporous ferns only into Eusporangiatae and Leptosporangiatae in 1884–9. The latter group was treated at a variety of ranks in subsequent systems of classification. The subclass "Polypodiidae" was first published and used for the homosporous leptosporangiate ferns by Cronquist, Takhtajan and Zimmermann in 1966, typified on Polypodium L.. Other contemporary classifications used the name "Filicidae" for this subclass. [10]
Smith et al. (2006) carried out the first higher-level classification of ferns based on molecular phylogenetics. They included heterosporous water ferns (Salviniales) (placed in a separate subclass by Cronquist et al. due to their highly modified morphology) within the leptosporangiate ferns, which they elevated to the rank of class as the Polypodiopsida (published by Cronquist et al. to include all ferns). [5]
The common ancestor of Salviniales, Cyatheales and Polypodiales went through a whole genome duplication. [11]
Later classifications renamed the group Polypodiidae, initially as a subclass of Equisetopsida sensu lato. [6] This subclass comprises leptosporangiate ferns as opposed to the remaining three subclasses which are informally referred to as eusporangiate ferns. The following diagram shows a likely phylogenic relationship between subclass Polypodiidae and the other Equisetopsida subclasses in that system [4]
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In 2014, Christenhusz and Chase grouped all the fern subclasses together as Polypodiophyta [3] and in 2016 the Pteridophyte Phylogeny Group (PPG) adopted the class Polypodiopsida sensu lato for the four fern subclasses. The following cladogram shows the phylogenic relationship between the subclasses according to the PPG. The first three small subclasses being informally grouped as eusporangiate ferns, in contrast to the Polypodiidae or leptosporangiate ferns. Polypodiidae is shown as a sister group of Marattiidae. [1]
Polypodiopsida |
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In both the Christenhusz and Chase, and the PPG classification, the extant Polypodiidae are divided into seven orders, 44 families, 300 genera, and an estimated 10,323 species. [1] [3]
Christenhusz and Chase 2014 [3] | Nitta et al. 2022 [12] and Fern Tree of life [13] | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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The following phylogram shows a likely relationship between the other vascular plant classes and the leptosporangiate ferns. It was formerly unclear about the relationship between Equisetopsida, Psilotopsida, and Marattiopsida, [14] [15] [16] but recent studies have shown that Equisetopsida is most likely sister to Psilotopsida.
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There has been some challenge to recent molecular studies, claiming that these provide a skewed view of the phylogenetic order because they do not take into account fossil representatives. [17] However, the molecular studies have clarified relations among families that had already been thought to be polyphyletic before the advent of molecular information but that were left in their polyphyletic ranks because there was not enough information to do otherwise. [18] The classification of ferns using these molecular studies, which have generally supported one another, reflects the best information available at present, because traditional morphological characters are not always informative in elucidating evolutionary relationships among ferns. [3]
The leptosporangiate ferns have a substantial fossil record. For example, fossils assigned to the Dicksoniaceae, a member of the Cyatheales, are known from the Lower Jurassic ( 201 to 175 million years ago). [19] A number of other extinct families have been described. They are not included in the classification systems used for extant ferns, and so most cannot be assigned to orders used in these systems. Taylor et al. (2009) use the order "Filicales", which corresponds to four Polypodiidae orders in more modern systems: Hymenophyllales, Gleicheniales, Schizaeales and Cyatheales. The unplaced families include: [20]
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.
The order Pinales in the division Pinophyta, class Pinopsida, comprises all the extant conifers. The distinguishing characteristic is the reproductive structure known as a cone produced by all Pinales. All of the extant conifers, such as Araucaria, cedar, celery-pine, cypress, fir, juniper, kauri, larch, pine, redwood, spruce, and yew, are included here. Some fossil conifers, however, belong to other distinct orders within the division Pinophyta.
Osmundaceae is a family of ferns containing four to six extant genera and 18–25 known species. It is the only living family of the order Osmundales in the class Polypodiopsida (ferns) or in some classifications the only order in the class Osmundopsida. This is an ancient and fairly isolated group that is often known as the "flowering ferns" because of the striking aspect of the ripe sporangia in Claytosmunda, Osmunda, Osmundastrum, and Plensium. In these genera the sporangia are borne naked on non-laminar pinnules, while Todea and Leptopteris bear sporangia naked on laminar pinnules. Ferns in this family are larger than most other ferns.
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.
Schizaeales is an order of ferns.
Equisetales is an order of subclass Equisetidae with only one living family, Equisetaceae, containing the genus Equisetum (horsetails).
The order Salviniales is an order of ferns in the class Polypodiopsida.
Marattiaceae is the only family of extant (living) ferns in the order Marattiales. In the Pteridophyte Phylogeny Group classification of 2016, 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.
Gleicheniales is an order of ferns in the subclass Polypodiidae. The Gleicheniales have records potentially as early as the Carboniferous, but the oldest unambiguous records date to the Permian.
The order Cyatheales, which includes most tree ferns, is a taxonomic order of the fern class, Polypodiopsida. No clear morphological features characterize all of the Cyatheales, but DNA sequence data indicate the order is monophyletic. Some species in the Cyatheales have tree-like growth forms from a vertical rhizome, others have shorter or horizontal expanding rhizomes.
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.
Equisetidae is one of the four subclasses of Polypodiopsida (ferns), a group of vascular plants with a fossil record going back to the Devonian. They are commonly known as horsetails. They typically grow in wet areas, with whorls of needle-like branches radiating at regular intervals from a single vertical stem.
A pteridophyte is a vascular plant that disperses 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.
The Hymenophyllaceae, the filmy ferns and bristle ferns, are a family of two to nine genera and about 650 known species of ferns, with a subcosmopolitan distribution, but generally restricted to very damp places or to locations where they are wetted by spray from waterfalls or springs. A recent fossil find shows that ferns of Hymenophyllaceae have existed since at least the Upper Triassic.
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.
Fern allies are a diverse group of seedless vascular plants that are not true ferns. Like ferns, a fern ally disperses by shedding spores to initiate an alternation of generations.
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.
The Pteridophyte Phylogeny Group (PPG) is an informal international group of systematic botanists who collaborate to establish a consensus on the classification of pteridophytes that reflects knowledge about plant relationships discovered through phylogenetic studies. In 2016, the group published a classification for extant pteridophytes, termed "PPG I". The paper had 94 authors.