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37-670: Genus: Species: Scoliosorus is a genus of ferns in the subfamily Vittarioideae of the family Pteridaceae with a single species, Scoliosorus ensiformis . The species is native to Mexico and Central America . This Pteridaceae -related article is a stub . You can help Misplaced Pages by expanding it . Fern The ferns ( Polypodiopsida or Polypodiophyta ) are a group of vascular plants (plants with xylem and phloem ) 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

74-734: A class Equisetopsida ( Embryophyta ) encompassing all land plants. This is referred to as Equisetopsida sensu lato to distinguish it from the narrower use to refer to horsetails alone, Equisetopsida sensu stricto . They placed the lycopods into subclass Lycopodiidae and the ferns, keeping the term monilophytes, into five subclasses, Equisetidae, Ophioglossidae, Psilotidae, Marattiidae and Polypodiidae, by dividing Smith's Psilotopsida into its two orders and elevating them to subclass (Ophioglossidae and Psilotidae). Christenhusz et al. (2011) followed this use of subclasses but recombined Smith's Psilotopsida as Ophioglossidae, giving four subclasses of ferns again. Christenhusz and Chase (2014) developed

111-634: A few species (e.g., Cyathea brownii on Norfolk Island and Cyathea medullaris in New Zealand ). Roots are underground non-photosynthetic structures that take up water and nutrients from soil . They are always fibrous and are structurally very similar to the roots of seed plants. As in all vascular plants , the sporophyte is the dominant phase or generation in the life cycle . The gametophytes of ferns, however, are very different from those of seed plants. They are free-living and resemble liverworts , whereas those of seed plants develop within

148-453: A frond is divided once into pinnae, the frond is called once pinnate. In some fronds the pinnae are further divided into segments, creating a bipinnate frond. The segments into which each pinna are divided are called pinnules, and the extensions of the rachis that support these pinnules, are called rachillae. Rarely, a frond may even be tripinnate, in which case the pinnule divisions are known as ultimate segments. Pinnae may be arranged along

185-487: A new classification of ferns and lycopods. They used the term Polypodiophyta for the ferns, subdivided like Smith et al. into four groups (shown with equivalents in the Smith system), with 21 families, approximately 212 genera and 10,535 species; This was a considerable reduction in the number of families from the 37 in the system of Smith et al., since the approach was more that of lumping rather than splitting. For instance

222-953: A number of families were reduced to subfamilies. Subsequently, a consensus group was formed, the Pteridophyte Phylogeny Group (PPG), analogous to the Angiosperm Phylogeny Group , publishing their first complete classification in November 2016. They recognise ferns as a class, the Polypodiopsida, with four subclasses as described by Christenhusz and Chase, and which are phylogenetically related as in this cladogram: Equisetales Ophioglossales Psilotales Marattiales Osmundales Hymenophyllales Gleicheniales Schizaeales Frond Fronds have particular terms describing their components. Like all leaves, fronds usually have

259-462: A protective coating called an indusium . The arrangement of the sporangia is important in classification. In monomorphic ferns, the fertile and sterile leaves look morphologically the same, and both are able to photosynthesize. In hemidimorphic ferns, just a portion of the fertile leaf is different from the sterile leaves. In dimorphic (holomorphic) ferns, the two types of leaves are morphologically distinct . The fertile leaves are much narrower than

296-430: A short-lived structure anchored to the ground by rhizoids called gametophyte which produce gametes. When a mature fertile frond bears sori, and spores are released, the spores will settle on the soil and send out rhizoids , while it develops into a prothallus . The prothallus bears spherical antheridia ( s.g. antheridium ) which produce antherozoids (male gametophytes) and archegonia ( s.g. archegonium ) which release

333-812: A significant input to the nitrogen nutrition of rice paddies . They also play certain roles in folklore. Extant ferns are herbaceous perennials and most lack woody growth. When woody growth is present, it is found in the stem. Their foliage may be deciduous or evergreen , and some are semi-evergreen depending on the climate. Like the sporophytes of seed plants, those of ferns consist of stems, leaves and roots. Ferns differ from spermatophytes in that they reproduce by spores rather than having flowers and producing seeds. However, they also differ from spore-producing bryophytes in that, like seed plants, they are polysporangiophytes , their sporophytes branching and producing many sporangia. Also unlike bryophytes, fern sporophytes are free-living and only briefly dependent on

370-568: A single oosphere . The antherozoid swims up the archegonium and fertilize the oosphere, resulting in a zygote, which will grow into a separate sporophyte, while the gametophyte shortly persists as a free-living plant. Carl Linnaeus (1753) originally recognized 15 genera of ferns and fern allies, classifying them in class Cryptogamia in two groups, Filices (e.g. Polypodium ) and Musci (mosses). By 1806 this had increased to 38 genera, and has progressively increased since ( see Schuettpelz et al (2018) ). Ferns were traditionally classified in

407-423: A stalk connecting them to the main stem. In botany , this leaf stalk is generally called a petiole , but in regard to fronds specifically it is called a stipe , and it supports a flattened blade (which may be called a lamina), and the continuation of the stipe into this portion is called the rachis . The blades may be simple (undivided), pinnatifid (deeply incised, but not truly compound), pinnate (compound with

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444-500: Is polyphyletic , the term fern allies should be abandoned, except in a historical context. More recent genetic studies demonstrated that the Lycopodiophyta are more distantly related to other vascular plants , having radiated evolutionarily at the base of the vascular plant clade , while both the whisk ferns and horsetails are as closely related to leptosporangiate ferns as the ophioglossoid ferns and Marattiaceae . In fact,

481-402: Is an outgrowth of the blade surface that may partly cover the sporangia. Some fern species feature frond dimorphism , in which fertile and sterile fronds differ in appearance and structure. Fern fronds, as with all leaves, arise from the stem, either directly, or on an outgrowth from the stem termed a phyllopodium . The stem of a typical (leptosporangiate) fern is subterranean or horizontal on

518-578: Is intermediate between the eusporangiate ferns and the leptosporangiate ferns. Rai and Graham (2010) broadly supported the primary groups, but queried their relationships, concluding that "at present perhaps the best that can be said about all relationships among the major lineages of monilophytes in current studies is that we do not understand them very well". Grewe et al. (2013) confirmed the inclusion of horsetails within ferns sensu lato , but also suggested that uncertainties remained in their precise placement. Other classifications have raised Ophioglossales to

555-780: Is usually called phyllody . The palm frond has been a symbol of victory, triumph, peace, and eternal life originating in the ancient Mediterranean world. For example, in some Christian traditions, during Palm Sunday, Jesus’ entrance into Jerusalem is celebrated by carrying palm leaves. During the Victorian phenomenon of Pteridomania or "fern craze", fern fronds became wildly popular symbols. Because fronds are somewhat flat, they could be used for decoration in ways that many other plants could not be. They were glued into collectors' albums, affixed to three dimensional objects, used as stencils for "spatter-work", inked and pressed into surfaces for nature printing, and so forth. The fern flower

592-646: The Cretaceous , contemporaneous with the rise of flowering plants that came to dominate the world's flora. Ferns are not of major economic importance, but some are used for food, medicine, as biofertilizer , as ornamental plants, and for remediating contaminated soil. They have been the subject of research for their ability to remove some chemical pollutants from the atmosphere. Some fern species, such as bracken ( Pteridium aquilinum ) and water fern ( Azolla filiculoides ), are significant weeds worldwide. Some fern genera, such as Azolla , can fix nitrogen and make

629-479: The class Filices, and later in a Division of the Plant Kingdom named Pteridophyta or Filicophyta. Pteridophyta is no longer recognised as a valid taxon because it is paraphyletic . The ferns are also referred to as Polypodiophyta or, when treated as a subdivision of Tracheophyta (vascular plants), Polypodiopsida, although this name sometimes only refers to leptosporangiate ferns. Traditionally, all of

666-485: The branched sporophyte is the dominant phase. Ferns have complex leaves called megaphylls that are more complex than the microphylls of clubmosses . Most ferns are leptosporangiate ferns . They produce coiled fiddleheads that uncoil and expand into fronds . The group includes about 10,560 known extant species. Ferns are defined here in the broad sense, being all of the Polypodiopsida , comprising both

703-429: The entire bodies of thalloid organisms, or the superficially leaf-like structures developed by some animals and fungi. Examples include frondose colonial bryozoans , extinct Ediacaran biota such as rangeomorphs , and some macroalgae and lichens. In paleontology of Ediacaran marine organisms, a frond may be defined as "a rangeomorph unit with a growth tip that can generate primary branches". A frond may also refer to

740-448: The entire frondose organism, including any stem or basal disc. To classify rangeomorph taxa, the frond is generally subdivided into segments as are those of a fern, and categorized by six factors: polarity, rows of branches, inflation, display/furling, alignment of branches, and presence of a basal disc. Frondescence is the production of leaves; it can also refer to the abnormal development of floral parts into leafy structures, though this

777-431: The fronds are branched more than once, it can also be a combination of the pinnatifid are pinnate shapes. If the leaf blades are divided twice, the plant has bipinnate fronds, and tripinnate fronds if they branch three times, and all the way to tetra- and pentapinnate fronds. In tree ferns, the main stalk that connects the leaf to the stem (known as the stipe), often has multiple leaflets. The leafy structures that grow from

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814-458: The group Ophioglossales have a unique arrangement -- such as a single fleshy or amorphous leaf. Fern fronds often bear sporangia , where the plant's spores are formed, usually on the underside (abaxial surface) of the pinnae, but sometimes marginally or scattered over the frond. The sporangia are typically clustered into a sorus (pl., sori). Associated with each sorus in many species is a membranous protective structure called an indusium, which

851-494: The inclusion of Equisetaceae in the ferns, notably relating to the construction of their sperm and peculiarities of their roots. The leptosporangiate ferns are sometimes called "true ferns". This group includes most plants familiarly known as ferns. Modern research supports older ideas based on morphology that the Osmundaceae diverged early in the evolutionary history of the leptosporangiate ferns; in certain ways this family

888-415: The leaflets arranged along a rachis to resemble a feather), or further compound (subdivided). If compound, a frond may be compound once, twice, or more. In a frond which is pinnate (feather-shaped), each leafy segment of the blade is called a pinna (plural pinnae), the stalk bearing the pinna is termed a petiolule, and the main vein or mid-rib of the pinna is referred to as a costa (plural costae). If

925-446: The leptosporangiate ( Polypodiidae ) and eusporangiate ferns , the latter group including horsetails , whisk ferns , marattioid ferns , and ophioglossoid ferns . The fern crown group , consisting of the leptosporangiates and eusporangiates, is estimated to have originated in the late Silurian period 423.2 million years ago, but Polypodiales , the group that makes up 80% of living fern diversity, did not appear and diversify until

962-474: The leptosporangiate ferns. The Marattiaceae are a primitive group of tropical ferns with large, fleshy rhizomes and are now thought to be a sibling taxon to the leptosporangiate ferns. Several other groups of species were considered fern allies: the clubmosses , spikemosses , and quillworts in Lycopodiophyta ; the whisk ferns of Psilotaceae ; and the horsetails of Equisetaceae . Since this grouping

999-555: The maternal gametophyte . The green , photosynthetic part of the plant is technically a megaphyll and in ferns, it is often called a frond . New leaves typically expand by the unrolling of a tight spiral called a crozier or fiddlehead into fronds . This uncurling of the leaf is termed circinate vernation . Leaves are divided into two types: sporophylls and tropophylls. Sporophylls produce spores; tropophylls do not. Fern spores are borne in sporangia which are usually clustered to form sori . The sporangia may be covered with

1036-544: The palm of a hand and have a short midrib or costa. Palmate fronds are also shaped like the palm of the hand, but all ribs or leaflets arise from a central area. A hastula is a flap of tissue borne at the insertion of the blade on the petiole on the upper, lower, or both leaf surfaces Bifurcate fronds may also develop. The extinct Devonian seed plant Cosmosperma polyloba demonstrated the early evolutionary diversification of frond branching patterns, presenting both bifurcate and trifurcate types. Some ferns, like members of

1073-462: The rachis either directly opposite one another or alternating up the stem. The arrangement may change from the base of a blade to the tip, as in the example of Blechnum shown below (from base to tip: pinnae opposite to alternate, and pinnatisect to pinnatifid). Some fronds are not pinnately compound (or simple), but may be palmate , costapalmate, or bifurcate . There is a spectrum from costapalmate to palmate. Costapalmate fronds are shaped like

1110-430: The rank of a fifth class, separating the whisk ferns and ophioglossoid ferns. The ferns are related to other groups as shown in the following cladogram: Lycophytes [REDACTED] Ferns [REDACTED] Gymnosperms [REDACTED] Angiosperms [REDACTED] The classification of Smith et al. in 2006 treated ferns as four classes: In addition they defined 11 orders and 37 families. That system

1147-476: The spore producing vascular plants were informally denominated the pteridophytes , rendering the term synonymous with ferns and fern allies . This can be confusing because members of the division Pteridophyta were also denominated pteridophytes ( sensu stricto ). Traditionally, three discrete groups have been denominated ferns: two groups of eusporangiate ferns, the families Ophioglossaceae ( adder's tongues , moonworts , and grape ferns) and Marattiaceae ; and

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1184-417: The spore wall and are dependent on the parent sporophyte for their nutrition. A fern gametophyte typically consists of: The lifecycle of a fern involves two stages, as in club mosses and horsetails . In stage one, the spores are produced by sporophytes in sporangia , which are clustered together in sori ( s.g. sorus ), developing on the underside of fertile fronds. In stage two, the spores germinate into

1221-496: The sterile leaves, and may have no green tissue at all, as in the Blechnaceae and Lomariopsidaceae . The anatomy of fern leaves can be anywhere from simple to highly divided, or even indeterminate (e.g. Gleicheniaceae , Lygodiaceae ). The divided forms are pinnate , where the leaf segments are completely separated from one other, or pinnatifid (partially pinnate), where the leaf segments are still partially connected. When

1258-463: The stipe are known as pinnae and are often again divided into smaller pinnules. Fern stems are often loosely called rhizomes , even though they grow underground only in some of the species. Epiphytic species and many of the terrestrial ones have above-ground creeping stolons (e.g., Polypodiaceae ), and many groups have above-ground erect semi-woody trunks (e.g., Cyatheaceae , the scaly tree ferns). These can reach up to 20 meters (66 ft) tall in

1295-423: The surface of the ground. These stems are called rhizomes . Many fern fronds are initially coiled into a fiddle-head or crozier (see circinate vernation ), although cycad and palm fronds do not have this pattern of new leaf growth. Fronds may bear hairs, scales, glands, and, in some species, bulblets for vegetative reproduction. Fronds may describe several " frondose " structures in non-plant organisms -- such as

1332-477: The whisk ferns and ophioglossoid ferns are demonstrably a clade , and the horsetails and Marattiaceae are arguably another clade. Smith et al. (2006) carried out the first higher-level pteridophyte classification published in the molecular phylogenetic era, and considered the ferns as monilophytes, as follows: Molecular data, which remain poorly constrained for many parts of the plants' phylogeny, have been supplemented by morphological observations supporting

1369-844: Was a consensus of a number of studies, and was further refined. The phylogenetic relationships are shown in the following cladogram (to the level of orders). This division into four major clades was then confirmed using morphology alone. Lycopodiophytes (club mosses, spike mosses, quillworts) Spermatophytes (seed plants) Equisetales (horsetails) [REDACTED] Ophioglossales (grapeferns etc.) Psilotales (whisk ferns) [REDACTED] Marattiales [REDACTED] Osmundales [REDACTED] Hymenophyllales (filmy ferns) [REDACTED] Gleicheniales [REDACTED] Schizaeales Salviniales (heterosporous) Cyatheales (tree ferns) [REDACTED] Polypodiales [REDACTED] Subsequently, Chase and Reveal considered both lycopods and ferns as subclasses of

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