Fern: Difference between revisions

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The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (land plants with vascular tissues such as xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from non-vascular plants (mosses, hornworts and liverworts) by having specialized transport bundles that conduct water and nutrients from and to the roots, as well as life cycles in which the branched sporophyte is the dominant phase.^[1]^[2]

Ferns have complex leaves called megaphylls that are more complex than the microphylls of clubmosses. Most ferns are leptosporangiate ferns that 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 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 during the rapid radiation of land plants,^[3] but Polypodiales, the group that makes up 80% of living fern diversity, did not appear and diversify until 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 a significant input to the nitrogen nutrition of rice paddies. They also play certain roles in folklore.

Description

Sporophyte

File:Samambaia (do tupi samambaîa) havaiana mini, em ambiente doméstico 02.jpg

Homegrown fern in Brazil

Extant ferns are herbaceous perennials and most lack woody growth.^[4] When woody growth is present, it is found in the stem.^[5] Their foliage may be deciduous or evergreen,^[6] and some are semi-evergreen depending on the climate.^[7] 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.^[5] 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 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.Template:Sfn 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 a protective coating called an indusium. The arrangement of the sporangia is important in classification.^[5]

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.^[8] The fertile leaves are much narrower than the sterile leaves, and may have no green tissue at all, as in the Blechnaceae and Lomariopsidaceae.

File:Croziers, fronds, rhizomes of bracken fern.jpg

Croziers, fronds, and rhizomes of bracken. In this species the stems grow underground, allowing the plant to spread horizontally.

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 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.^[9]^[10] 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 the stipe are known as pinnae and are often again divided into smaller pinnules.^[11]

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 Template:Convert tall in a few species (e.g., Cyathea brownii on Norfolk Island and Cyathea medullaris in New Zealand).^[12]

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.

Gametophyte

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 the spore wall and are dependent on the parent sporophyte for their nutrition.^[13] A fern gametophyte typically consists of:^[1]

Prothallus: A green, photosynthetic structure, whose initial growth is planar in one cell layer,^[14] usually heart or kidney shaped, 3–10 mm long and 2–8 mm broad.Script error: No such module "Unsubst". The prothallus produces gametes by means of:
- Antheridia: Small spherical structures that produce flagellate antherozoids.^[15]
- Archegonia: A flask-shaped structure that produces a single egg at the bottom, reached by the male gametophyte by swimming down the neck.^[1]
Rhizoids: root-like structures (not true roots) that consist of single^[16] greatly elongated cells, that absorb water and mineral salts over the whole structure. Rhizoids anchor the prothallus to the soil.^[1]

Life cycle and reproduction

File:Onoclea sensibilis 4 crop.jpg

New fern (Onoclea sensibilis) emerges from the prothallus.

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 short-lived gamete-producing structures called gametophytes anchored to the ground by rhizoids. When a mature fertile frond bears sori, and spores are released, the spores will settle on the soil and germinate to form initial rhizoids and protonemata that develop into the gametophyte's prothallus.^[17] The prothallus bears spherical antheridia (s.g. antheridium) which produce antherozoids (male gametophytes) and archegonia (s.g. archegonium) which release a single oosphere. The antherozoid swims up the archegonium and fertilizes the oosphere, resulting in a zygote, which will grow into a separate sporophyte, while the gametophyte shortly persists as a free-living plant.^[1]^[18]

Taxonomy

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).Template:Sfn Template:Sfn Template:Sfn By 1806 this had increased to 38 genera,Template:Sfn and has progressively increased since (see Template:Harvtxt). Ferns were traditionally classified in 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 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 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 is polyphyletic, the term fern allies should be abandoned, except in a historical context.Template:Sfn 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, the whisk ferns and ophioglossoid ferns are demonstrably a clade, and the horsetails and Marattiaceae are arguably another clade.

Molecular phylogenetics

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:Template:Sfn

Division Tracheophyta (tracheophytes) – vascular plants
- Sub division Euphyllophytina (euphyllophytes)
  - Infradivision Moniliformopses (monilophytes)
  - Infradivision Spermatophyta – seed plants, ~260,000 species
- Subdivision Lycopodiophyta (lycophytes) – less than 1% of extant vascular plants

Molecular data, which remain poorly constrained for many parts of the plants' phylogeny, have been supplemented by morphological observations supporting the inclusion of Equisetaceae in the ferns, notably relating to the construction of their sperm and peculiarities of their roots.Template:Sfn

The leptosporangiate ferns are sometimes called "true ferns".^[19] 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 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".^[20] Grewe et al. (2013) confirmed the inclusion of horsetails within ferns sensu lato, but also suggested that uncertainties remained in their precise placement.^[21] Other classifications have raised Ophioglossales to the rank of a fifth class, separating the whisk ferns and ophioglossoid ferns.^[21]

Phylogeny

The ferns are related to other groups as shown in the following cladogram:Template:Sfn Template:Sfn Template:Sfn Template:Sfn Template:Clade

Nomenclature and subdivision

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The classification of Smith et al. in 2006 treated ferns as four classes:Template:Sfn Template:Sfn

Equisetopsida (Sphenopsida) – 1 order, Equisetales (Horsetails) ~ 15 species
Psilotopsida – 2 orders (whisk ferns and ophioglossoid ferns) ~92 species
Marattiopsida – 1 order, Marattiales ~ 150 species
Polypodiopsida (Filicopsida) – 7 orders (leptosporangiate ferns) ~ 9,000 species

In addition they defined 11 orders and 37 families.Template:Sfn That system was a consensus of a number of studies, and was further refined.^[21]^[22] The phylogenetic relationships are shown in the following cladogram (to the level of orders).Template:Sfn^[23]^[21] This division into four major clades was then confirmed using morphology alone.Template:Sfn

Template:Fern cladogram.show(Orders & Outgroups)

Subsequently, Chase and Reveal considered both lycopods and ferns as subclasses of 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).Template:Sfn Christenhusz et al.Template:Efn (2011) followed this use of subclasses but recombined Smith's Psilotopsida as Ophioglossidae, giving four subclasses of ferns again.Template:Sfn

Christenhusz and Chase (2014) developed 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;Template:Sfn

Equisetidae (=Equisetopsida) – monotypic (Equisetales, Equisetaceae, Equisetum) horsetails ~ 20 species)
Ophioglossidae (=Psilotopsida) – 2 monotypic orders ~ 92 species
Marattiidae (=Marattiopsida) – 1 monotypic order (Marattiales, Marattiaceae, 2 subfamilies) ~ 130 species
Polypodiidae (=Polypodiopsida) – 7 orders

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 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 recognize ferns as a class, the Polypodiopsida, with four subclasses as described by Christenhusz and Chase, and which are phylogenetically related as in this cladogram:

Christenhusz and Chase 2014Template:Sfn	Nitta et al. 2022^[3] and Fern Tree of life^[24]
Template:Clade	Template:Clade

In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the Polypodiopsida consist of four subclasses, 11 orders, 48 families, 319 genera, and an estimated 10,578 species.Template:Sfn Thus Polypodiopsida in the broad sense (sensu lato) as used by the PPG (Polypodiopsida sensu PPG I) needs to be distinguished from the narrower usage (sensu stricto) of Smith et al. (Polypodiopsida sensu Smith et al.)Template:Sfn Classification of ferns remains unresolved and controversial with competing viewpoints (splitting vs lumping) between the systems of the PPG on the one hand and Christenhusz and Chase on the other, respectively. In 2018, Christenhusz and Chase explicitly argued against recognizing as many genera as PPG I.Template:Sfn Template:Sfn

Comparison of fern subdivisions in some classifications
Smith et al. (2006)Template:Sfn	Chase & Reveal (2009)Template:Sfn	Christenhusz et al. (2011)Template:Sfn	Christenhusz & Chase (2014, 2018)Template:Sfn Template:Sfn	PPG I (2016)Template:Sfn
ferns (no rank)	monilophytes (no rank)	ferns (monilophytes) (no rank)	ferns (Polypodiophyta) (no rank)	Class Polypodiopsida
Class Equisetopsida	Subclass Equisetidae	Subclass Equisetidae	Subclass Equisetidae	Subclass Equisetidae
Class Psilotopsida	Subclass Ophioglossidae Subclass Psilotidae	Subclass Ophioglossidae	Subclass Ophioglossidae	Subclass Ophioglossidae
Class Marattiopsida	Subclass Marattiidae	Subclass Marattiidae	Subclass Marattiidae	Subclass Marattiidae
Class Polypodiopsida	Subclass Polypodiidae	Subclass Polypodiidae	Subclass Polypodiidae	Subclass Polypodiidae

Evolution and biogeography

Fern-like taxa (Wattieza) first appear in the fossil record in the middle Devonian period, ca. 390 Mya. By the Triassic, the first evidence of ferns related to several modern families appeared. The great fern radiation occurred in the late Cretaceous, when many modern families of ferns first appeared.Template:Sfn Template:Sfn Template:Sfn Template:Sfn Ferns evolved to cope with low-light conditions present under the canopy of angiosperms.

Remarkably, the photoreceptor neochrome in the two orders Cyatheales and Polypodiales, integral to their adaptation to low-light conditions, was obtained via horizontal gene transfer from hornworts, a bryophyte lineage.^[25]

Due to the very large genome seen in most ferns, it was suspected they might have gone through whole genome duplications, but DNA sequencing has shown that their genome size is caused by the accumulation of mobile DNA like transposons and other genetic elements that infect genomes and get copied over and over again.^[26]

Ferns appear to have evolved extrafloral nectaries 135 million years ago, nearly simultaneously with the trait's evolution in angiosperms. However, nectary-associated diversifications in ferns did not hit their stride until nearly 100 million years later, in the Cenozoic. There is weak support for the rise of fern-feeding arthropods driving this diversification.^[27]

Distribution and habitat

Ferns are widespread in their distribution, with the greatest richness in the tropics and least in arctic areas. The greatest diversity occurs in tropical rainforests.Template:Sfn New Zealand, for which the fern is a symbol, has about 230 species, distributed throughout the country.Template:Sfn It is a common plant in European forests.

Ecology

Fern species live in a wide variety of habitats, from remote mountain elevations, to dry desert rock faces, bodies of water or open fields. Ferns in general may be thought of as largely being specialists in marginal habitats, often succeeding in places where various environmental factors limit the success of flowering plants. Some ferns are among the world's most serious weed species, including the bracken fern growing in the Scottish highlands, or the mosquito fern (Azolla) growing in tropical lakes, both species forming large aggressively spreading colonies. There are four particular types of habitats that ferns are found in: moist, shady forests; crevices in rock faces, especially when sheltered from the full sun; acid wetlands including bogs and swamps; and tropical trees, where many species are epiphytes (something like a quarter to a third of all fern species).Template:Sfn

Especially the epiphytic ferns have turned out to be hosts of a huge diversity of invertebrates. It is assumed that bird's-nest ferns alone contain up to half the invertebrate biomass within a hectare of rainforest canopy.^[28]

Many ferns depend on associations with mycorrhizal fungi. Many ferns grow only within specific pH ranges; for instance, the climbing fern (Lygodium palmatum) of eastern North America will grow only in moist, intensely acid soils, while the bulblet bladder fern (Cystopteris bulbifera), with an overlapping range, is found only on limestone.

The spores are rich in lipids, protein and calories, so some vertebrates eat these. The European woodmouse (Apodemus sylvaticus) has been found to eat the spores of Culcita macrocarpa, and the bullfinch (Pyrrhula murina) and the New Zealand lesser short-tailed bat (Mystacina tuberculata) also eat fern spores.^[29]

In undergrowth below coast redwoods, California

In undergrowth below coast redwoods, California
Fern bed under a forest canopy, Virginia

Fern bed under a forest canopy, Virginia
On a wall

On a wall
Asplenium hart's tongue fern in a gryke in limestone pavement

Asplenium hart's tongue fern in a gryke in limestone pavement
Epiphytic ferns in India

Epiphytic ferns in India
Azolla duckweed fern covering the Canning River, Western Australia

Azolla duckweed fern covering
the Canning River, Western Australia

Life cycle

Ferns are vascular plants differing from lycophytes by having true leaves (megaphylls), which are often pinnate. They differ from seed plants (gymnosperms and angiosperms) in reproducing by means of spores and lacking flowers and seeds. Like all land plants, they have a life cycle referred to as alternation of generations, characterized by alternating diploid sporophytic and haploid gametophytic phases. The diploid sporophyte has 2n paired chromosomes, where n varies from species to species. The haploid gametophyte has n unpaired chromosomes, i.e. half the number of the sporophyte. The gametophyte of ferns is a free-living organism, whereas the gametophyte of the gymnosperms and angiosperms is dependent on the sporophyte.

The life cycle of a typical fern proceeds as follows:

A diploid sporophyte phase produces haploid spores by meiosis (a process of cell division which reduces the number of chromosomes by a half).
A spore grows into a free-living haploid gametophyte by mitosis (a process of cell division which maintains the number of chromosomes). The gametophyte typically consists of a photosynthetic prothallus.
The gametophyte produces gametes (often both sperm and eggs on the same prothallus) by mitosis.
A mobile, flagellate sperm fertilizes an egg that remains attached to the prothallus.
The fertilized egg is now a diploid zygote and grows by mitosis into a diploid sporophyte (the typical fern plant).

Sometimes a gametophyte can give rise to sporophyte traits like roots or sporangia without the rest of the sporophyte.^[30]

Sorus of monarch fern, with sporangium

Sorus of monarch fern, with sporangium
Gametophyte (thallus) and sporophyte (ascendant frond) of Onoclea sensibilis

Gametophyte (thallus) and sporophyte (ascendant frond) of Onoclea sensibilis

Uses

Ferns are not as important economically as seed plants, but have considerable importance in some societies. Some ferns are used for food, including the fiddleheads of Pteridium aquilinum (bracken), Matteuccia struthiopteris (ostrich fern), and Osmundastrum cinnamomeum (cinnamon fern). Diplazium esculentum is also used in the tropics (for example in budu pakis, a traditional dish of Brunei^[31]) as food. Tubers from the "para", Ptisana salicina (king fern) are a traditional food in New Zealand and the South Pacific. Fern tubers were used for food 30,000 years ago in Europe.^[32]^[33] Fern tubers were used by the Guanches to make gofio in the Canary Islands. Ferns are generally not known to be poisonous to humans.^[34] Licorice fern rhizomes were chewed by the natives of the Pacific Northwest for their flavor.^[35] Some species of ferns are carcinogenic, and the British Royal Horticultural Society has advised not to consume any species for health reasons of both humans and livestock.^[36]

Ferns of the genus Azolla, commonly known as water fern or mosquito ferns are very small, floating plants that do not resemble ferns. The mosquito ferns are used as a biological fertilizer in the rice paddies of southeast Asia, taking advantage of their ability to fix nitrogen from the air into compounds that can then be used by other plants.

Ferns have proved resistant to phytophagous insects. The gene that express the protein Tma12 in an edible fern, Tectaria macrodonta, has been transferred to cotton plants, which became resistant to whitefly infestations.^[37]

Many ferns are grown in horticulture as landscape plants, for cut foliage and as houseplants, especially the Boston fern (Nephrolepis exaltata) and other members of the genus Nephrolepis. The bird's nest fern (Asplenium nidus) is also popular, as are the staghorn ferns (genus Platycerium). Perennial (also known as hardy) ferns planted in gardens in the northern hemisphere also have a considerable following.^[38]

Several ferns, such as bracken^[39] and Azolla^[40] species are noxious weeds or invasive species. Further examples include Japanese climbing fern (Lygodium japonicum), sensitive fern (Onoclea sensibilis) and Giant water fern (Salvinia molesta), one of the world's worst aquatic weeds.^[41]^[42] The important fossil fuel coal consists of the remains of primitive plants, including ferns.^[43]

Culture

File:Nature print, Alois Auer.jpg

Ferns in the Victorian era: Blätter des Manns Walfarn by Alois Auer, Vienna: Imperial Printing Office, 1853

Pteridology

The study of ferns and other pteridophytes is called pteridology. A pteridologist is a specialist in the study of pteridophytes in a broader sense that includes the more distantly related lycophytes.

Pteridomania

Pteridomania was a Victorian era craze which involved fern collecting and fern motifs in decorative art including pottery, glass, metals, textiles, wood, printed paper, and sculpture "appearing on everything from christening presents to gravestones and memorials." The fashion for growing ferns indoors led to the development of the Wardian case, a glazed cabinet that would exclude air pollutants and maintain the necessary humidity.^[44]

Other applications

File:Barnsley fern plotted with VisSim.PNG

Barnsley fern created using a chaos game, through an Iterated function system^[45]

The Barnsley fern is a fractal named after the British mathematician Michael Barnsley who first described it in his book Fractals Everywhere. A self-similar structure is described by a mathematical function, applied repeatedly at different scales to create a frond pattern.^[45]

The dried form of ferns was used in other arts, such as a stencil or directly inked for use in a design. The botanical work, The Ferns of Great Britain and Ireland, is a notable example of this type of nature printing. The process, patented by the artist and publisher Henry Bradbury, impressed a specimen on to a soft lead plate. The first publication to demonstrate this was Alois Auer's The Discovery of the Nature Printing-Process.

Fern bars were popular in America in the 1970s and 80s.

Folklore

Ferns figure in folklore, for example in legends about mythical flowers or seeds.Template:Sfn In Slavic folklore, ferns are believed to bloom once a year, during the Ivan Kupala night. Although alleged to be exceedingly difficult to find, anyone who sees a fern flower is thought to be guaranteed to be happy and rich for the rest of their life. Similarly, Finnish tradition holds that one who finds the seed of a fern in bloom on Midsummer night will, by possession of it, be guided and be able to travel invisibly to the locations where eternally blazing Will o' the wisps called aarnivalkea mark the spot of hidden treasure. These spots are protected by a spell that prevents anyone but the fern-seed holder from ever knowing their locations.^[46] In Wicca, ferns are thought to have magical properties such as a dried fern can be thrown into hot coals of a fire to exorcise evil spirits, or smoke from a burning fern is thought to drive away snakes and such creatures.^[47]

New Zealand

The Silver fern is the national emblem of New Zealand and feature on its passport and in the design of its national airline, Air New Zealand, and of its rugby team, the All Blacks.Script error: No such module "Unsubst".

Organisms confused with ferns

Misnomers

Several non-fern plants (and even animals) are called ferns and are sometimes confused with ferns. These include:

Asparagus fern—This may apply to one of several species of the monocot genus Asparagus, which are flowering plants.
Sweetfern—A flowering shrub of the genus Comptonia.
Air fern—A group of animals called hydrozoans that are distantly related to jellyfish and corals. They are harvested, dried, dyed green, and then sold as a plant that can live on air. While it may look like a fern, it is merely the skeleton of this colonial animal.
Fern bush—Chamaebatiaria millefolium—a rose family shrub with fern-like leaves.
Fern tree—Jacaranda mimosifolia—an ornamental tree of the order Lamiales.
Fern leaf tree—Filicium decipiens—an ornamental tree of the order Sapindales.

Fern-like flowering plants

Some flowering plants such as palms and members of the carrot family have pinnate leaves that somewhat resemble fern fronds. However, these plants have fully developed seeds contained in fruits, rather than the microscopic spores of ferns.

Notes

Template:Notelist

References

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Bibliography

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Books

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Lord, Thomas R. (2006). Ferns and Fern Allies of Pennsylvania. Indiana, Pennsylvania: Pinelands Press. Ferns and Fern Allies of Pennsylvania – Thomas Reeves Lord.
Moran, Robbin C. (2004). A Natural History of Ferns. Portland, Oregon: Timber Press. Template:ISBN
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Journal articles

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Websites

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A classification of the ferns and their allies. (Australian National Herbarium)
A fern book bibliography.
Register of fossil Pteridophyta
Watson, L. and M. J. Dallwitz (2004 onwards). The Ferns (Filicopsida) of the British Isles. Template:Webarchive
Ferns and Pteridomania in Victorian Scotland.
Non-seed plant images at bioimages.vanderbilt.edu
American Fern Society
British Pteridological Society Template:Webarchive
International Equisetological Association

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Fern: Difference between revisions

Latest revision as of 04:42, 20 November 2025

Contents

Description

Sporophyte

Gametophyte

Life cycle and reproduction

Taxonomy

Molecular phylogenetics

Phylogeny

Nomenclature and subdivision

Evolution and biogeography

Distribution and habitat

Ecology

Life cycle

Uses

Culture

Pteridology

Pteridomania

Other applications

Folklore

New Zealand

Organisms confused with ferns

Misnomers

Fern-like flowering plants

See also

Notes

References

Bibliography

Books

Journal articles

Websites

Navigation menu

@@ Line 12: / Line 12: @@
 |caption1        = [[Psilotales]]
 |image2          = Equisetopsida.jpg
 |caption2        = [[Equisetales]]
 |image3          = Flickr - brewbooks - Angiopteris evecta - Mule's foot fern (1).jpg
 |caption3        = [[Marattiales]]
@@ Line 45: / Line 45: @@
 }}
-The '''ferns''' ('''Polypodiopsida''' or '''Polypodiophyta''') are a group of [[vascular plant]]s (plants with [[xylem]] and [[phloem]]) that reproduce via [[spore]]s and have neither [[seed]]s nor [[flower]]s. They differ from [[moss]]es 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.<ref name=":0">{{Cite book |title=The Ultimate Family Visual Dictionary |publisher=[[DK Pub.]] |year=2012 |isbn=978-0-1434-1954-9 |location=New Delhi |pages=118-121 |language=en}}</ref>
+The '''ferns''' ('''Polypodiopsida''' or '''Polypodiophyta''') are a group of [[vascular plant]]s ([[land plant]]s with [[vascular tissue]]s such as [[xylem]] and [[phloem]]) that [[plant reproduction|reproduce]] via [[spore]]s and have neither [[seed]]s nor [[flower]]s. They differ from [[non-vascular plant]]s ([[moss]]es, [[hornwort]]s and [[liverwort]]s) by having specialized [[vascular bundle|transport bundle]]s that conduct [[water]] and [[nutrient]]s from and to the [[root]]s, as well as [[biological life cycle|life cycle]]s in which the branched [[sporophyte]] is the dominant phase.<ref name="DK Pub.-2017">{{Cite book|edition=11th|title=Ultimate Visual Dictionary|publisher=[[DK Pub.]]|year=2017|isbn=978-1-4654-5894-0|url=|pages=120–121|language=en|chapter=Horsetails, clubmosses, and ferns|chapter-url=https://archive.org/details/ultimate-visual-dictionary-by-dk-z-lib.org/page/120/mode/2up|via=Internet Archive}}</ref><ref>{{cite journal|title=Early evolution of life cycles in embryophytes: A focus on the fossil evidence of gametophyte/sporophyte size and morphological complexity|last1=Gerrienne|first1=Philippe|last2=Gonez|first2=Paul|journal=Journal of Systematics and Evolution|year=2011|volume=49|issue=1|pages=1-16|doi=10.1111/j.1759-6831.2010.00096.x|doi-access=free}}</ref>
-Ferns have complex [[leaf|leaves]] called [[megaphyll]]s that are more complex than the [[microphyll]]s of [[clubmosses]]. Most ferns are [[leptosporangiate fern]]s. They produce coiled [[Fiddlehead fern|fiddleheads]] that uncoil and expand into [[frond]]s. The group includes about 10,560 known extant species. Ferns are defined here in the broad sense, being all of the [[Polypodiopsida]], comprising both the leptosporangiate ([[Polypodiidae (plant)|Polypodiidae]]) and [[eusporangiate fern]]s, the latter group including [[horsetail]]s, [[Psilotaceae|whisk ferns]], [[marattioid fern]]s, and [[ophioglossoid fern]]s.
+Ferns have complex [[leaf|leaves]] called [[megaphyll]]s that are more complex than the [[microphyll]]s of [[clubmoss]]es. Most ferns are [[leptosporangiate fern]]s that produce coiled [[Fiddlehead fern|fiddleheads]] that uncoil and expand into [[frond]]s. The group includes about 10,560 known extant species. Ferns are defined here in the broad sense, being all of the [[Polypodiopsida]], comprising both the leptosporangiate ([[Polypodiidae (plant)|Polypodiidae]]) and [[eusporangiate fern]]s, the latter group including [[horsetail]]s, [[Psilotaceae|whisk ferns]], [[marattioid fern]]s and [[ophioglossoid fern]]s.
-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,<ref name="Polypodiopsida">{{cite journal |last1=Nitta |first1=Joel H. |last2=Schuettpelz |first2=Eric |last3=Ramírez-Barahona |first3=Santiago |last4=Iwasaki |first4=Wataru |display-authors=et al. |year=2022 |title=An Open and Continuously Updated Fern Tree of Life |journal=Frontiers in Plant Science |volume=13 |page=909768 |doi=10.3389/fpls.2022.909768 |pmid=36092417 |pmc=9449725 |doi-access=free}}</ref> but [[Polypodiales]], the group that makes up 80% of living fern diversity, did not appear and diversify until the [[Cretaceous]], contemporaneous with the rise of flowering plants that came to dominate the world's flora.
+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 during the [[Silurian-Devonian Terrestrial Revolution|rapid radiation of land plants]],<ref name="Polypodiopsida">{{cite journal |last1=Nitta |first1=Joel H. |last2=Schuettpelz |first2=Eric |last3=Ramírez-Barahona |first3=Santiago |last4=Iwasaki |first4=Wataru |display-authors=et al. |year=2022 |title=An Open and Continuously Updated Fern Tree of Life |journal=Frontiers in Plant Science |volume=13 |article-number=909768 |doi=10.3389/fpls.2022.909768 |pmid=36092417 |pmc=9449725 |doi-access=free|bibcode=2022FrPS...1309768N }}</ref> but [[Polypodiales]], the group that makes up 80% of living fern diversity, did not appear and diversify until the [[Cretaceous]], contemporaneous with the [[Cretaceous Terrestrial Revolution|rise of]] [[flowering plant]]s 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 [[nitrogen fixation|fix nitrogen]] and make a significant input to the nitrogen nutrition of [[Paddy field|rice paddies]]. They also play certain roles in folklore.
+Ferns are not of major economic importance, but some are used for [[food]], [[medicinal plants|medicine]], as [[biofertilizer]], as [[ornamental plant]]s, 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 [[weed]]s worldwide. Some fern genera, such as ''[[Azolla]]'', can [[nitrogen fixation|fix nitrogen]] and make a significant input to the nitrogen nutrition of [[paddy field|rice paddies]]. They also play certain roles in folklore.
 ==Description==
 === Sporophyte ===
+[[Image:Samambaia (do tupi samambaîa) havaiana mini, em ambiente doméstico 02.jpg|thumb|Homegrown fern in Brazil]]
 Extant ferns are herbaceous [[Perennial plant|perennials]] and most lack [[wood]]y growth.<ref>{{Cite book |last=Mauseth |first=James D. |url=https://books.google.com/books?id=UEsBPPlxP7EC&pg=PA492 |title=Botany: an Introduction to Plant Biology |date=September 2008 |publisher=Jones & Bartlett Publishers |isbn=978-1-4496-4720-9 |page=492}}</ref> When woody growth is present, it is found in the stem.<ref name="Levyns-1966">{{Cite book |last=Levyns |first=M. R. |title=A Guide to the Flora of the Cape Peninsula |publisher=Juta & Company |year=1966 |edition=2nd Revised |oclc=621340}}</ref> Their foliage may be [[deciduous]] or [[evergreen]],<ref>{{Cite book |last1=Fernández |first1=Helena |url=https://books.google.com/books?id=smtpBM7UgXEC&pg=PA175 |title=Working with Ferns: Issues and Applications |last2=Kumar |first2=Ashwani |last3=Revilla |first3=Maria Angeles |date=2010-11-11 |publisher=Springer |isbn=978-1-4419-7162-3 |page=175}}</ref> and some are semi-evergreen depending on the climate.<ref>{{Cite book |last=Hodgson |first=Larry |url=https://books.google.com/books?id=HDij6Fmk2MwC&pg=PA329 |title=Making the Most of Shade: How to Plan, Plant, and Grow a Fabulous Garden that Lightens Up the Shadows |date=2005-01-01 |publisher=Rodale |isbn=978-1-57954-966-4 |page=329}}</ref> Like the sporophytes of seed plants, those of ferns consist of stems, leaves and roots. Ferns differ from [[spermatophyte]]s in that they reproduce by spores rather than having flowers and producing seeds.<ref name="Levyns-1966" /> However, they also differ from spore-producing [[bryophyte]]s in that, like seed plants, they are [[polysporangiophyte]]s, their [[sporophyte]]s branching and producing many sporangia. Also unlike bryophytes, fern sporophytes are free-living and only briefly dependent on the maternal [[gametophyte]].
-The [[green]], [[photosynthesis|photosynthetic]] part of the plant is technically a [[microphyll#evolution of leaves: microphyll and megaphyll|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 fern|fiddlehead]] into [[frond]]s.{{sfn|McCausland|2019}} This uncurling of the leaf is termed [[circinate vernation]]. Leaves are divided into two types: sporophylls and tropophylls. [[Sporophyll]]s produce spores; [[tropophyll]]s do not. Fern spores are borne in [[sporangia]] which are usually clustered to form [[sorus|sori]]. The sporangia may be covered with a protective coating called an [[indusium]]. The arrangement of the sporangia is important in classification.<ref name="Levyns-1966" />
+The [[green]], [[photosynthesis|photosynthetic]] part of the plant is technically a [[microphyll#evolution of leaves: microphyll and megaphyll|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 fern|fiddlehead]] into [[frond]]s.{{sfn|McCausland|2019}} This uncurling of the leaf is termed [[circinate vernation]]. Leaves are divided into two types: sporophylls and tropophylls. [[Sporophyll]]s produce spores; [[tropophyll]]s do not. Fern spores are borne in [[sporangia]] which are usually clustered to form [[sorus|sori]]. The sporangia may be covered with a protective coating called an [[indusium]]. The arrangement of the [[Sporangium|sporangia]] is important in classification.<ref name="Levyns-1966" />
-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 [[frond dimorphism|morphologically distinct]].<ref>[https://2019.botanyconference.org/engine/search/index.php?func=detail&aid=758 Understanding the contribution of LFY and PEBP flowering genes to fern leaf dimorphism – Botany 2019].</ref> The fertile leaves are much narrower than the sterile leaves, and may have no green tissue at all, as in the [[Blechnaceae]] and [[Lomariopsidaceae]].
+In monomorphic ferns, the fertile and sterile leaves look morphologically the same, and both are able to [[Photosynthesis|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 [[frond dimorphism|morphologically distinct]].<ref>[https://2019.botanyconference.org/engine/search/index.php?func=detail&aid=758 Understanding the contribution of LFY and PEBP flowering genes to fern leaf dimorphism – Botany 2019].</ref> The fertile leaves are much narrower than the sterile leaves, and may have no green tissue at all, as in the [[Blechnaceae]] and [[Lomariopsidaceae]].
 [[File:Croziers, fronds, rhizomes of bracken fern.jpg|thumb|Croziers, fronds, and [[rhizome]]s of bracken. In this species the stems grow underground, allowing the plant to spread horizontally.]]
-The anatomy of fern leaves can be anywhere from simple to highly divided, or even [[Indeterminate growth|indeterminate]] (e.g. [[Gleicheniaceae]], [[Lygodium|Lygodiaceae]]). The divided forms are [[Pinnation|pinnate]], where the leaf segments are completely separated from one other, or pinnatifid (partially pinnate), where the leaf segments are still partially connected. When 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.<ref>[https://www.fs.fed.us/wildflowers/beauty/ferns/structure.shtml Fern Structure – Forest Service].</ref><ref>[https://www.nzplants.auckland.ac.nz/en/about/ferns/structure/frond.html Fern Structure – Forest Service], Auckland, New Zealand.</ref> 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 the stipe are known as pinnae and are often again divided into smaller pinnules.<ref>{{cite web |url=http://www.basicbiology.net/plants/ferns/fronds.php |title=Fern Fronds |publisher=Basic Biology |access-date=2014-12-06 |archive-date=19 April 2015 |archive-url=https://web.archive.org/web/20150419003216/http://basicbiology.net/plants/ferns/fronds.php |url-status=dead}}</ref>
+The anatomy of fern leaves can be anywhere from simple to highly divided, or even [[Indeterminate growth|indeterminate]] (e.g. [[Gleicheniaceae]], [[Lygodium|Lygodiaceae]]). The divided forms are [[Pinnation|pinnate]], where the leaf segments are completely separated from one other, or pinnatifid (partially pinnate), where the leaf segments are still partially connected. When the fronds are branched more than once, it can also be a combination of the pinnatifid are [[Pinnation|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.<ref>[https://www.fs.fed.us/wildflowers/beauty/ferns/structure.shtml Fern Structure – Forest Service].</ref><ref>[https://www.nzplants.auckland.ac.nz/en/about/ferns/structure/frond.html Fern Structure – Forest Service], Auckland, New Zealand.</ref> 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 the stipe are known as pinnae and are often again divided into smaller pinnules.<ref>{{cite web |url=http://www.basicbiology.net/plants/ferns/fronds.php |title=Fern Fronds |publisher=Basic Biology |access-date=2014-12-06 |archive-date=19 April 2015 |archive-url=https://web.archive.org/web/20150419003216/http://basicbiology.net/plants/ferns/fronds.php }}</ref>
-Fern stems are often loosely called [[rhizome]]s, even though they grow underground only in some of the species. Epiphytic species and many of the terrestrial ones have above-ground creeping [[stolon]]s (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 {{convert|20|m|ft|sp=us}} tall in a few species (e.g., ''[[Cyathea brownii]]'' on [[Norfolk Island]] and ''[[Cyathea medullaris]]'' in [[New Zealand]]).<ref>{{cite book |first1=Mark F. |last1=Large |first2=John E. |last2=Braggins |author-link2=John E. Braggins |date=2004 |title=Tree Ferns |publisher=Timber Press |isbn=0881926302 |url-access=registration |url=https://archive.org/details/treeferns00mark |page= }}</ref>
+Fern stems are often loosely called [[rhizome]]s, even though they grow underground only in some of the species. Epiphytic species and many of the terrestrial ones have above-ground creeping [[stolon]]s (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 {{convert|20|m|ft|sp=us}} tall in a few species (e.g., ''[[Cyathea brownii]]'' on [[Norfolk Island]] and ''[[Cyathea medullaris]]'' in [[New Zealand]]).<ref>{{cite book |first1=Mark F. |last1=Large |first2=John E. |last2=Braggins |author-link2=John E. Braggins |date=2004 |title=Tree Ferns |publisher=Timber Press |isbn=0-88192-630-2 |url-access=registration |url=https://archive.org/details/treeferns00mark |page= }}</ref>
-Roots are underground non-photosynthetic structures that take up water and nutrients from [[soil]]. They are always [[fibrous root system|fibrous]] and are structurally very similar to the roots of seed plants.
+[[Root|Roots]] are underground non-photosynthetic structures that take up water and nutrients from [[soil]]. They are always [[fibrous root system|fibrous]] and are structurally very similar to the roots of seed plants.
 === Gametophyte ===
-As in all [[vascular plant]]s, the sporophyte is the dominant phase or [[alternation of generations|generation in the life cycle]]. The [[gametophyte]]s 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 the spore wall and are dependent on the parent sporophyte for their nutrition. A fern gametophyte typically consists of:<ref name=":0" />
+As in all [[vascular plant]]s, the [[sporophyte]] is the dominant phase or [[alternation of generations|generation in the life cycle]]. The [[gametophyte]]s 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 the spore wall and are dependent on the parent sporophyte for their nutrition.<ref>{{cite journal|last1=Villareal A.|first1=Juan Carlos|last2=Campos S.|first2=Laura Victoria|last3=Urbide-M.|first3=Jaime|last4=Goffinet|first4=Bernard|title=Parallel Evolution of Endospory within Hornworts: ''Nothoceros renzagliensis'' (Dendrocerotaceae), sp. nov.|journal=Systematic Botany|volume=37|issue=1|pages=31-37|doi=10.1600/036364412X616594}}</ref> A fern gametophyte typically consists of:<ref name="DK Pub.-2017" />
+* [[Prothallus]]: A green, photosynthetic structure, whose initial growth is planar in one cell layer,<ref>{{cite journal|last1=Kelley|first1=A.G.|last2=Postlethwait|first2=S.N.|title=Fern Gametophytes as a Tool for the Study of Morphogenesis|pages=56-60|year=1960|volume=70|journal=Proceedings of the Indiana Academy of Science|url=https://journals.indianapolis.iu.edu/index.php/ias/article/view/5459|access-date=16 November 2025}}</ref> usually heart or kidney shaped, 3–10 mm long and 2–8 mm broad.{{citation needed|date=November 2025}} The prothallus produces gametes by means of:
-* [[Prothallus]]: A green, photosynthetic structure that is one cell thick, usually heart or kidney shaped, 3–10 mm long and 2–8 mm broad. The prothallus produces gametes by means of:
+** [[Antheridium|Antheridia]]: Small spherical structures that produce [[flagellum|flagellate]] antherozoids.<ref>{{cite encyclopedia|title=A Dictionary of Plant Sciences|edition=2nd|editor-last=Allaby|editor-first=Michael|entry-url=https://www.oxfordreference.com/display/10.1093/acref/9780198608912.001.0001/acref-9780198608912-e-386|entry=antherozoid|isbn=9780191726804|publisher=Oxford University Press|year=2006}}</ref>
-** [[Antheridium|Antheridia]]: Small spherical structures that produce [[flagellum|flagellate]] antherozoids.<ref name=":0" />
+** [[Archegonium|Archegonia]]: A flask-shaped structure that produces a single egg at the bottom, reached by the male gametophyte by swimming down the neck.<ref name="DK Pub.-2017" />
-** [[Archegonium|Archegonia]]: A flask-shaped structure that produces a single egg at the bottom, reached by the male gametophyte by swimming down the neck.<ref name=":0" />
+* [[Rhizoid]]s: [[root]]-like structures (not true roots) that consist of single<ref>{{cite journal|title=The evolution of root hairs and rhizoids|last1=Jones|first1=Victor A.S.|last2=Dolan|first2=Liam|journal=Annals of Botany|volume=110|issue=2|pages=205-212|doi=10.1093/aob/mcs136|pmc=3394659|pmid=22730024|year=2012}}</ref> greatly elongated cells, that absorb water and mineral [[salts]] over the whole structure. Rhizoids anchor the prothallus to the soil.<ref name="DK Pub.-2017" />
-* [[Rhizoid]]s: [[root]]-like structures (not true roots) that consist of single greatly elongated cells, that absorb water and mineral [[salts]] over the whole structure. Rhizoids anchor the prothallus to the soil.<ref name=":0" />
 == Life cycle and reproduction ==
 [[File:Onoclea sensibilis 4 crop.jpg|thumb|New fern (''[[Onoclea sensibilis]]'') emerges from the [[prothallus]].]]
-The lifecycle of a fern involves two stages, as in [[Lycopodiopsida|club mosses]] and [[Equisetum|horsetails]]. In stage one, the spores are produced by [[Sporophyte|sporophytes]] in [[Sporangium|sporangia]], which are clustered together in sori (''s.g. sorus''), developing on the underside of fertile fronds. In stage two, the spores germinate into 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 [[Rhizoid|rhizoids]], while it develops into a [[prothallus]]. The prothallus bears spherical [[Antheridium|antheridia]] (''s.g. antheridium'') which produce antherozoids (male gametophytes) and [[Archegonium|archegonia]] (''s.g. archegonium'') which release a single [[Egg cell|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.<ref name=":0" />
+The lifecycle of a fern involves two stages, as in [[Lycopodiopsida|club mosses]] and [[Equisetum|horsetails]]. In stage one, the spores are produced by [[sporophyte]]s in [[Sporangium|sporangia]], which are clustered together in sori ({{abbr|s.g.|singular}} ''sorus''), developing on the underside of fertile fronds. In stage two, the spores germinate into short-lived gamete-producing structures called [[gametophyte]]s anchored to the ground by [[rhizoid]]s. When a mature fertile frond bears sori, and spores are released, the spores will settle on the soil and germinate to form initial rhizoids and protonemata that develop into the gametophyte's [[prothallus]].<ref>{{cite journal|last=Racusen|first=Richard H.|title=Early Development in Fern Gametophytes: Interpreting the Transition to Prothallial Architecture in Terms of Coordinated Photosynthate Production and Osmotic Ion Uptake|volume=89|issue=2|journal=Annals of Botany|doi=10.1093/aob/mcf032|pmid=12099354|pmc=4233796|year=2002}}</ref>  The prothallus bears spherical [[Antheridium|antheridia]] (s.g. ''antheridium'') which produce antherozoids (male gametophytes) and [[Archegonium|archegonia]] (s.g. ''archegonium'') which release a single [[Egg cell|oosphere]]. The antherozoid swims up the archegonium and fertilizes the oosphere, resulting in a zygote, which will grow into a separate sporophyte, while the gametophyte shortly persists as a free-living plant.<ref name="DK Pub.-2017" /><ref>{{cite journal|title=Providing the missing links in fern life history: Insights from a phenological survey of the gametophyte stage|journal=Applications in Plant Sciences|last1=Quinlain|first1=Alexandra|last2=Lee|first2=Pei-Hsuan|last3=Tang|first3=Te-Yen|last4=Huang|first4=Yao-Moan|last5=Chiou|first5=Wen-Liang|last6=Kuo|first6=Li-Yaung|volume=10|issue=2|article-number=e11473|year=2022|doi=10.1002/aps3.11473|pmc=9039788|pmid=35495188|doi-access=free}}</ref>
 == Taxonomy ==
@@ Line 101: / Line 99: @@
 Molecular data, which remain poorly constrained for many parts of the plants' phylogeny, have been supplemented by morphological observations supporting the inclusion of Equisetaceae in the ferns, notably relating to the construction of their sperm and peculiarities of their roots.{{sfn|Smith et al.2006}}
-The leptosporangiate ferns are sometimes called "true ferns".<ref>{{Cite book |last=Stace |first=Clive |author-link = Clive Stace |year=2010b |title=New Flora of the British Isles |edition=3rd |location=Cambridge, UK |publisher=Cambridge University Press |isbn=978-0-521-70772-5 |page=xxviii}}</ref> 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 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".<ref name=RaiGraham2010>{{Cite journal |last1=Rai |first1=Hardeep S. |last2=Graham |first2=Sean W. |year=2010 |title=Utility of a large, multigene plastid data set in inferring higher-order relationships in ferns and relatives (monilophytes) |journal=American Journal of Botany |volume=97 |issue=9 |pages=1444–1456 |doi=10.3732/ajb.0900305 |name-list-style=amp |pmid=21616899}}, p. 1450</ref> Grewe et al. (2013) confirmed the inclusion of horsetails within ferns ''sensu lato'', but also suggested that uncertainties remained in their precise placement.<ref name=Grew13/> Other classifications have raised Ophioglossales to the rank of a fifth class, separating the whisk ferns and ophioglossoid ferns.<ref name=Grew13/>
+The leptosporangiate ferns are sometimes called "true ferns".<ref>{{Cite book |last=Stace |first=Clive |author-link = Clive Stace |year=2010b |title=New Flora of the British Isles |edition=3rd |location=Cambridge, UK |publisher=Cambridge University Press |isbn=978-0-521-70772-5 |page=xxviii}}</ref> 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 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".<ref name=RaiGraham2010>{{Cite journal |last1=Rai |first1=Hardeep S. |last2=Graham |first2=Sean W. |year=2010 |title=Utility of a large, multigene plastid data set in inferring higher-order relationships in ferns and relatives (monilophytes) |journal=American Journal of Botany |volume=97 |issue=9 |pages=1444–1456 |doi=10.3732/ajb.0900305 |name-list-style=amp |pmid=21616899|bibcode=2010AmJB...97.1444R }}, p. 1450</ref> Grewe et al. (2013) confirmed the inclusion of horsetails within ferns ''sensu lato'', but also suggested that uncertainties remained in their precise placement.<ref name=Grew13/> Other classifications have raised Ophioglossales to the rank of a fifth class, separating the whisk ferns and ophioglossoid ferns.<ref name=Grew13/>
 === Phylogeny ===
@@ Line 107: / Line 105: @@
 The ferns are related to other groups as shown in the following cladogram:{{sfn|Christenhusz|Chase|2014}}{{sfn|Cantino et al|2007}}{{sfn|Chase|Reveal|2009}}{{sfn|Pteridophyte Phylogeny Group|2016}}
 {{clade |style=line-height:140%;font-size:100%;
 |label1= [[Tracheophyta]] |sublabel1= (vascular plants)
 |1={{clade
-    |1= [[Lycopodiophyta|Lycophytes]] [[File:2021-03_Amsterdam_Island_-_lycopodium_clavatum_10.jpg|50px]]
+    |1= [[Lycopodiophyta|Lycophytes]] [[File:2021-03 Amsterdam Island - lycopodium clavatum 10.jpg|50px]]
     |label2= [[Euphyllophyta]]
     |2={{clade
-       |1='''Ferns''' [[File:California_Arena_Point_fern.jpg|70px]]
+       |1='''Ferns''' [[File:California Arena Point fern.jpg|70px]]
        |label2= [[Spermatophyta]] |sublabel2=(seed plants)
        |2={{clade
-          |1=[[Gymnosperm]]s [[File:Flore_des_serres_v15_147a.jpg|50px]]
+          |1=[[Gymnosperm]]s [[File:Flore des serres v15 147a.jpg|50px]]
           |2=[[Angiosperm]]s [[File:160 Ranunculus repens.jpg|50px]]
           }}
@@ Line 127: / Line 125: @@
 The classification of Smith et al. in 2006 treated ferns as four classes:{{sfn|Smith et al.2006}}{{sfn|Schuettpelz|2007|loc=Table I}}
 * [[Equisetidae|Equisetopsida]] (Sphenopsida) – 1 order, [[Equisetales]] ([[Horsetail]]s) ~ 15 species
 * [[Ophioglossidae|Psilotopsida]] – 2 orders ([[Psilotaceae|whisk ferns]] and [[ophioglossoid fern]]s) ~92 species
 * [[Marattiidae|Marattiopsida]] – 1 order, [[Marattiales]] ~ 150 species
 * [[Polypodiidae (plant)|Polypodiopsida]] (Filicopsida) – 7 orders ([[leptosporangiate fern]]s) ~ 9,000 species
-In addition they defined 11 orders and 37 families.{{sfn|Smith et al.2006}} That system was a consensus of a number of studies, and was further refined.<ref name=Grew13>{{cite journal |last=Grewe |first=Felix |title=Complete plastid genomes from Ophioglossum californicum, Psilotum nudum, and Equisetum hyemale reveal an ancestral land plant genome structure and resolve the position of Equisetales among monilophytes |journal=BMC Evolutionary Biology |year=2013 |volume=13 |issue=1 |pages=1–16 |doi=10.1186/1471-2148-13-8 |issn=1471-2148 |display-authors=etal |pmid=23311954 |pmc=3553075 |doi-access=free |bibcode=2013BMCEE..13....8G }}</ref><ref>{{cite journal |last=Karol |first=Kenneth G. |title=Complete plastome sequences of Equisetum arvense and Isoetes flaccida: implications for phylogeny and plastid genome evolution of early land plant lineages |journal=BMC Evolutionary Biology |year=2010|volume=10 |pages=321–336 |doi=10.1186/1471-2148-10-321 |issn=1471-2148 |pmid=20969798|pmc=3087542|issue=1|display-authors=etal |doi-access=free |bibcode=2010BMCEE..10..321K }}</ref> The phylogenetic relationships are shown in the following [[cladogram]] (to the level of orders).{{sfn|Smith et al.2006}}<ref>{{cite journal |last1=Li |first1=F-W |last2=Kuo |first2=L-Y |last3=Rothfels |first3=CJ |last4=Ebihara |first4=A |last5=Chiou |first5=W-L |display-authors=etal |year=2011 |title=rbcL and matK Earn Two Thumbs Up as the Core DNA Barcode for Ferns |journal=PLOS ONE |volume=6 |issue=10|page=e26597 |doi=10.1371/journal.pone.0026597 |pmid=22028918 |pmc=3197659|bibcode=2011PLoSO...626597L |doi-access=free }}</ref><ref name=Grew13/> This division into four major clades was then confirmed using [[plant morphology|morphology]] alone.{{sfn|Schneider et al|2009}}
+In addition they defined 11 orders and 37 families.{{sfn|Smith et al.2006}} That system was a consensus of a number of studies, and was further refined.<ref name=Grew13>{{cite journal |last=Grewe |first=Felix |title=Complete plastid genomes from Ophioglossum californicum, Psilotum nudum, and Equisetum hyemale reveal an ancestral land plant genome structure and resolve the position of Equisetales among monilophytes |journal=BMC Evolutionary Biology |year=2013 |volume=13 |issue=1 |pages=1–16 |doi=10.1186/1471-2148-13-8 |issn=1471-2148 |display-authors=etal |article-number=8 |pmid=23311954 |pmc=3553075 |doi-access=free |bibcode=2013BMCEE..13....8G }}</ref><ref>{{cite journal |last=Karol |first=Kenneth G. |title=Complete plastome sequences of Equisetum arvense and Isoetes flaccida: implications for phylogeny and plastid genome evolution of early land plant lineages |journal=BMC Evolutionary Biology |year=2010|volume=10 |pages=321–336 |doi=10.1186/1471-2148-10-321 |issn=1471-2148 |pmid=20969798|pmc=3087542|issue=1|display-authors=etal |doi-access=free |bibcode=2010BMCEE..10..321K }}</ref> The phylogenetic relationships are shown in the following [[cladogram]] (to the level of orders).{{sfn|Smith et al.2006}}<ref>{{cite journal |last1=Li |first1=F-W |last2=Kuo |first2=L-Y |last3=Rothfels |first3=CJ |last4=Ebihara |first4=A |last5=Chiou |first5=W-L |display-authors=etal |year=2011 |title=rbcL and matK Earn Two Thumbs Up as the Core DNA Barcode for Ferns |journal=PLOS ONE |volume=6 |issue=10|article-number=e26597 |doi=10.1371/journal.pone.0026597 |pmid=22028918 |pmc=3197659|bibcode=2011PLoSO...626597L |doi-access=free }}</ref><ref name=Grew13/> This division into four major clades was then confirmed using [[plant morphology|morphology]] alone.{{sfn|Schneider et al|2009}}
 {{Fern cladogram.show(Orders & Outgroups)}}
@@ Line 145: / Line 142: @@
 * [[Polypodiidae (plant)|Polypodiidae]] (=Polypodiopsida) – 7 orders
-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 a number of families were reduced to subfamilies. Subsequently, a [[Scientific consensus|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:
+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 a number of families were reduced to subfamilies. Subsequently, a [[Scientific consensus|consensus]] group was formed, the [[Pteridophyte Phylogeny Group]] (PPG), analogous to the [[Angiosperm Phylogeny Group]], publishing their first complete classification in November 2016. They recognize ferns as a class, the Polypodiopsida, with four subclasses as described by Christenhusz and Chase, and which are phylogenetically related as in this cladogram:
 {|class="wikitable"
@@ Line 151: / Line 148: @@
 ! colspan=1 |Christenhusz and Chase 2014{{sfn|Pteridophyte Phylogeny Group|2016}}
 ! colspan=1 |Nitta et al. 2022<ref name="Polypodiopsida"/> and Fern Tree of life<ref>{{cite web |year=2024 |title=Tree viewer: interactive visualization of FTOL |url=https://fernphy.github.io/viewer.html |version=FTOL v1.7.0 |access-date=1 May 2025}}</ref>
 |-
 |style="vertical-align:top|
 {{Clade |style=font-size:90%;line-height:80%
@@ Line 241: / Line 238: @@
 ! Smith et al. (2006){{sfn|Smith et al.2006}} !! Chase & Reveal (2009){{sfn|Chase|Reveal|2009}} !! Christenhusz et al. (2011){{sfn|Christenhusz et al|2011}} !! Christenhusz & Chase (2014, 2018){{sfn|Christenhusz|Chase|2014}}{{sfn|Christenhusz et al|2018}} !! PPG I (2016){{sfn|Pteridophyte Phylogeny Group|2016}}
 |-
-|'''ferns'''<br/>(no rank) ||'''monilophytes'''<br/>(no rank) ||'''ferns (monilophytes)'''<br/>(no rank) ||'''ferns (Polypodiophyta)'''<br/>(no rank) ||&nbsp;'''Class Polypodiopsida'''
+|'''ferns'''<br />(no rank) ||'''monilophytes'''<br />(no rank) ||'''ferns (monilophytes)'''<br />(no rank) ||'''ferns (Polypodiophyta)'''<br />(no rank) ||&nbsp;'''Class Polypodiopsida'''
 |-
 |Class Equisetopsida ||&nbsp;&nbsp;Subclass Equisetidae ||&nbsp;&nbsp;Subclass Equisetidae ||&nbsp;&nbsp;Subclass Equisetidae ||&nbsp;Subclass [[Equisetidae]]
 |-
-|Class Psilotopsida ||&nbsp;&nbsp;Subclass Ophioglossidae<br/>&nbsp;&nbsp;Subclass Psilotidae ||&nbsp;&nbsp;Subclass Ophioglossidae ||&nbsp;&nbsp;Subclass Ophioglossidae ||&nbsp;Subclass [[Ophioglossidae]]
+|Class Psilotopsida ||&nbsp;&nbsp;Subclass Ophioglossidae<br />&nbsp;&nbsp;Subclass Psilotidae ||&nbsp;&nbsp;Subclass Ophioglossidae ||&nbsp;&nbsp;Subclass Ophioglossidae ||&nbsp;Subclass [[Ophioglossidae]]
 |-
 |Class Marattiopsida ||&nbsp;&nbsp;Subclass Marattiidae ||&nbsp;&nbsp;Subclass Marattiidae ||&nbsp;&nbsp;Subclass Marattiidae ||&nbsp;Subclass [[Marattiidae]]
@@ Line 260: / Line 257: @@
 Due to the very large genome seen in most ferns, it was suspected they might have gone through [[Paleopolyploidy|whole genome duplications]], but [[DNA sequencing]] has shown that their genome size is caused by the accumulation of mobile DNA like [[Transposable element|transposons]] and other genetic elements that infect genomes and get copied over and over again.<ref>[https://www.science.org/content/article/genes-seeds-arose-early-plant-evolution-ferns-reveal Genes for seeds arose early in plant evolution, ferns reveal]</ref>
-Ferns appear to have evolved [[extrafloral nectaries]] 135 million years ago, nearly simultaneously with the trait's evolution in angiosperms. However, nectary-associated diversifications in ferns did not hit their stride until nearly 100 million years later, in the [[Cenozoic]]. There is weak support for the rise of fern-feeding arthropods driving this diversification.<ref>{{Cite journal |last1=Suissa |first1=Jacob S. |last2=Li |first2=Fay-Wei |last3=Moreau |first3=Corrie S. |date=2024-05-24 |title=Convergent evolution of fern nectaries facilitated independent recruitment of ant-bodyguards from flowering plants |journal=Nature Communications |language=en |volume=15 |issue=1 |pages=4392 |doi=10.1038/s41467-024-48646-x |issn=2041-1723|doi-access=free |pmid=38789437 |pmc=11126701 |bibcode=2024NatCo..15.4392S }}</ref>
+Ferns appear to have evolved [[extrafloral nectaries]] 135 million years ago, nearly simultaneously with the trait's evolution in angiosperms. However, nectary-associated diversifications in ferns did not hit their stride until nearly 100 million years later, in the [[Cenozoic]]. There is weak support for the rise of fern-feeding arthropods driving this diversification.<ref>{{Cite journal |last1=Suissa |first1=Jacob S. |last2=Li |first2=Fay-Wei |last3=Moreau |first3=Corrie S. |date=2024-05-24 |title=Convergent evolution of fern nectaries facilitated independent recruitment of ant-bodyguards from flowering plants |journal=Nature Communications |language=en |volume=15 |issue=1 |page=4392 |doi=10.1038/s41467-024-48646-x |issn=2041-1723|doi-access=free |pmid=38789437 |pmc=11126701 |bibcode=2024NatCo..15.4392S }}</ref>
 == Distribution and habitat ==
@@ Line 268: / Line 265: @@
 == Ecology ==
-Fern species live in a wide variety of [[habitat]]s, from remote [[mountain]] elevations, to dry [[desert]] rock faces, bodies of water or open fields. Ferns in general may be thought of as largely being specialists in marginal habitats, often succeeding in places where various environmental factors limit the success of [[flowering plant]]s. Some ferns are among the world's most serious weed species, including the [[bracken]] fern growing in the Scottish highlands, or the mosquito fern (''[[Azolla]]'') growing in tropical lakes, both species forming large aggressively spreading colonies. There are four particular types of habitats that ferns are found in: moist, shady [[forest]]s; crevices in rock faces, especially when sheltered from the full sun; acid wetlands including [[bog]]s and [[swamp]]s; and tropical [[tree]]s, where many species are [[epiphyte]]s (something like a quarter to a third of all fern species).{{sfn|Schuettpelz|2007|loc=Part I}}
+Fern species live in a wide variety of [[habitat]]s, from remote [[mountain]] elevations, to dry [[desert]] rock faces, bodies of water or open fields. Ferns in general may be thought of as largely being specialists in marginal habitats, often succeeding in places where various environmental factors limit the success of [[flowering plant]]s. Some ferns are among the world's most serious [[weed]] species, including the [[bracken]] fern growing in the Scottish highlands, or the mosquito fern (''[[Azolla]]'') growing in tropical lakes, both species forming large aggressively spreading colonies. There are four particular types of habitats that ferns are found in: moist, shady [[forest]]s; crevices in rock faces, especially when sheltered from the full sun; acid wetlands including [[bog]]s and [[swamp]]s; and tropical [[tree]]s, where many species are [[epiphyte]]s (something like a quarter to a third of all fern species).{{sfn|Schuettpelz|2007|loc=Part I}}
 Especially the epiphytic ferns have turned out to be hosts of a huge diversity of invertebrates. It is assumed that [[bird's-nest fern]]s alone contain up to half the invertebrate biomass within a hectare of [[rainforest]] canopy.<ref>{{Cite web|url=https://www.science.org/content/article/ferns-brimming-life|title=Ferns Brimming With Life|date=2 June 2004|website=Science &#124; AAAS}}</ref>
@@ Line 274: / Line 271: @@
 Many ferns depend on associations with [[mycorrhizal]] fungi. Many ferns grow only within specific pH ranges; for instance, the climbing fern (''[[Lygodium palmatum]]'') of eastern [[North America]] will grow only in moist, intensely [[acid]] soils, while the bulblet bladder fern (''[[Cystopteris bulbifera]]''), with an overlapping range, is found only on [[limestone]].
-The spores are rich in [[lipid]]s, [[protein]] and [[calories]], so some vertebrates eat these. The [[European woodmouse]] (''Apodemus sylvaticus'') has been found to eat the spores of ''[[Culcita macrocarpa]]'', and the [[bullfinch]] (''Pyrrhula murina'') and the [[New Zealand lesser short-tailed bat]] (''Mystacina tuberculata'') also eat fern spores.<ref>{{cite news|url=http://news.bbc.co.uk/earth/hi/earth_news/newsid_8523000/8523825.stm|title=A mouse that eats ferns like a dinosaur|last=Walker|first=Matt|date=19 February 2010|publisher=BBC Earth News|access-date=2010-02-20}}</ref>
+The spores are rich in [[lipid]]s, [[protein]] and [[calories]], so some vertebrates eat these. The [[European woodmouse]] (''Apodemus sylvaticus'') has been found to eat the spores of ''[[Culcita macrocarpa]]'', and the [[bullfinch]] (''Pyrrhula murina'') and the [[New Zealand lesser short-tailed bat]] (''Mystacina tuberculata'') also eat fern spores.<ref>{{cite news|url=https://news.bbc.co.uk/earth/hi/earth_news/newsid_8523000/8523825.stm|title=A mouse that eats ferns like a dinosaur|last=Walker|first=Matt|date=19 February 2010|publisher=BBC Earth News|access-date=2010-02-20}}</ref>
 <gallery class=center mode=nolines widths=300 heights=250>
@@ Line 282: / Line 279: @@
 File:Harts Tongue Fern on the Great Orme - geograph.org.uk - 345785.jpg|''[[Asplenium]]'' hart's tongue fern in a [[wikt:gryke|gryke]] in [[limestone pavement]]
 File:... epiphytic fern (49095803342).jpg|[[Epiphyte|Epiphytic]] ferns in India
-File:Canning_rv_azolla_10_gnangarra.jpg|''[[Azolla]]'' duckweed fern covering<br/>the [[Canning River]], Western Australia
+File:Canning_rv_azolla_10_gnangarra.jpg|''[[Azolla]]'' duckweed fern covering<br />the [[Canning River]], Western Australia
 </gallery>
@@ Line 305: / Line 302: @@
 == Uses ==
-Ferns are not as important economically as seed plants, but have considerable importance in some societies. Some ferns are used for food, including the fiddleheads of ''Pteridium aquilinum'' ([[bracken]]), ''Matteuccia struthiopteris'' ([[ostrich fern]]), and ''Osmundastrum cinnamomeum'' ([[cinnamon fern]]). ''[[Diplazium esculentum]]'' is also used in the tropics (for example in ''budu pakis'', a traditional dish of [[Brunei]]<ref>{{cite book|title=Indigenous Fermented Foods of Southeast Asia|year=2015}}</ref>) as food. Tubers from the "para", ''[[Ptisana salicina]]'' (king fern) are a traditional food in [[New Zealand]] and the [[Oceania|South Pacific]]. Fern tubers were used for food 30,000 years ago in Europe.<ref>Van Gilder Cooke, Sonia (23 October 2010). [https://www.newscientist.com/article/dn19597-stone-age-humans-liked-their-burgers-in-a-bun.html "Stone Age humans liked their burgers in a bun".] ''[[New Scientist]]'', p. 18.</ref><ref>Revedin, Anna et al. (18 October 2010). [http://www.pnas.org/cgi/doi/10.1073/pnas.1006993107 "Thirty thousand-year-old evidence of plant food processing".] ''[[PNAS]]''.</ref> Fern tubers were used by the [[Guanches]] to make [[gofio]] in the [[Canary Islands]]. Ferns are generally not known to be poisonous to humans.<ref name="edible plant">{{cite book | title=The Official Pocket Edible Plant Survival Manual | publisher=Freedom and Liberty Foundation Press | last=Pelton |first=Robert | year=2011 | page=25 | id=[[Barnes & Noble|BNID]] [http://www.barnesandnoble.com/w/?ean=2940013382145 2940013382145]}}</ref> [[Licorice fern]] [[rhizome]]s were chewed by the natives of the [[Pacific Northwest]] for their flavor.<ref>{{Cite book |last=Moerman |first=Daniel E. |url=https://books.google.com/books?id=iYhjlKR7GZEC |title=Native American Food Plants: An Ethnobotanical Dictionary |date=2010-10-27 |publisher=Timber Press |isbn=978-1-60469-189-4 |pages=190 |language=en}}</ref> Some species of ferns are [[carcinogenic]], and the British Royal Horticultural Society has advised not to consume any species for health reasons of both humans and livestock.<ref>{{cite web|title=Dol Sot Bibimbap|url=http://apps.rhs.org.uk/advicesearch/Profile.aspx?pid=445|access-date=2011-12-19|archive-url=https://web.archive.org/web/20111111190952/http://apps.rhs.org.uk/advicesearch/profile.aspx?PID=445|archive-date=2011-11-11|url-status=dead}}</ref>
+Ferns are not as important economically as seed plants, but have considerable importance in some societies. Some ferns are used for food, including the fiddleheads of ''Pteridium aquilinum'' ([[bracken]]), ''Matteuccia struthiopteris'' ([[ostrich fern]]), and ''Osmundastrum cinnamomeum'' ([[cinnamon fern]]). ''[[Diplazium esculentum]]'' is also used in the tropics (for example in ''budu pakis'', a traditional dish of [[Brunei]]<ref>{{cite book|title=Indigenous Fermented Foods of Southeast Asia|year=2015}}</ref>) as food. Tubers from the "para", ''[[Ptisana salicina]]'' (king fern) are a traditional food in [[New Zealand]] and the [[Oceania|South Pacific]]. Fern tubers were used for food 30,000 years ago in Europe.<ref>Van Gilder Cooke, Sonia (23 October 2010). [https://www.newscientist.com/article/dn19597-stone-age-humans-liked-their-burgers-in-a-bun.html "Stone Age humans liked their burgers in a bun".] ''[[New Scientist]]'', p. 18.</ref><ref>Revedin, Anna et al. (18 October 2010). [http://www.pnas.org/cgi/doi/10.1073/pnas.1006993107 "Thirty thousand-year-old evidence of plant food processing".] ''[[PNAS]]''.</ref> Fern tubers were used by the [[Guanches]] to make [[gofio]] in the [[Canary Islands]]. Ferns are generally not known to be poisonous to humans.<ref name="edible plant">{{cite book | title=The Official Pocket Edible Plant Survival Manual | publisher=Freedom and Liberty Foundation Press | last=Pelton |first=Robert | year=2011 | page=25 | id=[[Barnes & Noble|BNID]] [http://www.barnesandnoble.com/w/?ean=2940013382145 2940013382145]}}</ref> [[Licorice fern]] [[rhizome]]s were chewed by the natives of the [[Pacific Northwest]] for their flavor.<ref>{{Cite book |last=Moerman |first=Daniel E. |url=https://books.google.com/books?id=iYhjlKR7GZEC |title=Native American Food Plants: An Ethnobotanical Dictionary |date=2010-10-27 |publisher=Timber Press |isbn=978-1-60469-189-4 |page=190 |language=en}}</ref> Some species of ferns are [[carcinogenic]], and the British Royal Horticultural Society has advised not to consume any species for health reasons of both humans and livestock.<ref>{{cite web|title=Dol Sot Bibimbap|url=http://apps.rhs.org.uk/advicesearch/Profile.aspx?pid=445|access-date=2011-12-19|archive-url=https://web.archive.org/web/20111111190952/http://apps.rhs.org.uk/advicesearch/profile.aspx?PID=445|archive-date=2011-11-11}}</ref>
 Ferns of the genus ''Azolla'', commonly known as water fern or mosquito ferns are very small, floating plants that do not resemble ferns. The mosquito ferns are used as a biological fertilizer in the rice paddies of southeast Asia, taking advantage of their ability to [[nitrogen fixation|fix nitrogen]] from the air into compounds that can then be used by other plants.
@@ Line 313: / Line 310: @@
 Many ferns are grown in [[horticulture]] as landscape plants, for [[foliage|cut foliage]] and as [[houseplant]]s, especially the Boston fern (''[[Nephrolepis exaltata]]'') and other members of the genus ''[[Nephrolepis]]''. The [[Asplenium nidus|bird's nest fern]] (''Asplenium nidus'') is also popular, as are the [[staghorn fern]]s (genus ''Platycerium''). Perennial (also known as hardy) ferns planted in gardens in the northern hemisphere also have a considerable following.<ref>{{Cite web |title=Ferns: A Classic Shade Garden Plant |url=https://extension.sdstate.edu/ferns-classic-shade-garden-plant |access-date=2023-05-30 |website=extension.sdstate.edu}}</ref>
-Several ferns, such as bracken<ref name=CABIPteridium>{{cite web|date=2018 |title=Datasheet: ''Pteridium aquilinum'' (bracken) |url=https://www.cabi.org/isc/datasheet/45596 |publisher=CAB International |access-date=11 February 2019 }}</ref> and ''Azolla''<ref name=CABIAzolla>{{cite web|date=2018 |title=Datasheet: ''Azolla filiculoides'' (water fern) |url=https://www.cabi.org/isc/datasheet/8119 |publisher=CAB International |access-date=11 February 2019 }}</ref> species are noxious [[weed]]s or [[invasive species]]. Further examples include Japanese climbing fern (''[[Lygodium japonicum]]''), [[sensitive fern]] (''Onoclea sensibilis'') and Giant water fern (''[[Salvinia molesta]]''), one of the world's worst aquatic weeds.<ref>{{Cite web |title={{!}} Center for Aquatic and Invasive Plants {{!}} University of Florida, IFAS |url=https://plants.ifas.ufl.edu/plant-directory/salvinia-molesta/ |access-date=2023-05-30 |website=plants.ifas.ufl.edu}}</ref><ref>{{Cite book|last=Moran|first=Robbin|title=A Natural History of Ferns|year=2004|publisher=Timber Press |isbn=0-88192-667-1}}</ref> The important fossil fuel [[coal]] consists of the remains of primitive plants, including ferns.<ref>{{Cite web |title=Fossils, Kentucky Geological Survey, University of Kentucky |url=https://www.uky.edu/KGS/fossils/fossil-pennsylvanianPlants.php |access-date=2023-05-30 |website=www.uky.edu}}</ref>
+Several ferns, such as bracken<ref name=CABIPteridium>{{cite web|date=2018 |title=Datasheet: ''Pteridium aquilinum'' (bracken) |url=https://www.cabi.org/isc/datasheet/45596 |publisher=CAB International |access-date=11 February 2019 }}</ref> and ''Azolla''<ref name=CABIAzolla>{{cite web|date=2018 |title=Datasheet: ''Azolla filiculoides'' (water fern) |url=https://www.cabi.org/isc/datasheet/8119 |publisher=CAB International |access-date=11 February 2019 }}</ref> species are noxious [[weed]]s or [[invasive species]]. Further examples include Japanese climbing fern (''[[Lygodium japonicum]]''), [[sensitive fern]] (''Onoclea sensibilis'') and Giant water fern (''[[Salvinia molesta]]''), one of the world's worst aquatic weeds.<ref>{{Cite web |title={{!}} Center for Aquatic and Invasive Plants {{!}} University of Florida, IFAS |url=https://plants.ifas.ufl.edu/plant-directory/salvinia-molesta/ |access-date=2023-05-30 |website=plants.ifas.ufl.edu |archive-date=30 May 2023 |archive-url=https://web.archive.org/web/20230530232110/https://plants.ifas.ufl.edu/plant-directory/salvinia-molesta/ }}</ref><ref>{{Cite book|last=Moran|first=Robbin|title=A Natural History of Ferns|year=2004|publisher=Timber Press |isbn=0-88192-667-1}}</ref> The important fossil fuel [[coal]] consists of the remains of primitive plants, including ferns.<ref>{{Cite web |title=Fossils, Kentucky Geological Survey, University of Kentucky |url=https://www.uky.edu/KGS/fossils/fossil-pennsylvanianPlants.php |access-date=2023-05-30 |website=www.uky.edu}}</ref>
 == Culture ==
@@ Line 339: / Line 336: @@
 === Folklore ===
-Ferns figure in folklore, for example in legends about mythical flowers or seeds.{{sfn|May|1978}} In [[Slavic folklore]], ferns are believed to bloom once a year, during the [[Ivan Kupala Day|Ivan Kupala]] night. Although alleged to be exceedingly difficult to find, anyone who sees a [[fern flower]] is thought to be guaranteed to be happy and rich for the rest of their life. Similarly, [[Finland|Finnish]] tradition holds that one who finds the [[seed]] of a fern in bloom on [[Midsummer]] night will, by possession of it, be guided and be able to travel invisibly to the locations where eternally blazing [[Will o' the wisp]]s called ''[[aarnivalkea]]'' mark the spot of hidden [[treasure]]. These spots are protected by a spell that prevents anyone but the fern-seed holder from ever knowing their locations.<ref>{{cite web |url=http://www.saunalahti.fi/~marian1/gourmet/season5a.htm |title=Traditional Finnish Midsummer celebration |publisher=Saunalahti.fi |access-date=2013-09-07 |archive-date=19 September 2011 |archive-url=https://web.archive.org/web/20110919201014/http://www.saunalahti.fi/~marian1/gourmet/season5a.htm |url-status=dead }}</ref> In Wicca, ferns are thought to have magical properties such as a dried fern can be thrown into hot coals of a fire to exorcise evil spirits, or smoke from a burning fern is thought to drive away snakes and such creatures.<ref name="Cunningham">{{cite book |last=Cunningham |first=Scott |author-link=Scott Cunningham |title=Cunningham's Encyclopedia of Magical Herbs |date=1999 |publisher=Llewellyn |page=102}}</ref>
+Ferns figure in folklore, for example in legends about mythical flowers or seeds.{{sfn|May|1978}} In [[Slavic folklore]], ferns are believed to bloom once a year, during the [[Ivan Kupala Day|Ivan Kupala]] night. Although alleged to be exceedingly difficult to find, anyone who sees a [[fern flower]] is thought to be guaranteed to be happy and rich for the rest of their life. Similarly, [[Finland|Finnish]] tradition holds that one who finds the [[seed]] of a fern in bloom on [[Midsummer]] night will, by possession of it, be guided and be able to travel invisibly to the locations where eternally blazing [[Will o' the wisp]]s called ''[[aarnivalkea]]'' mark the spot of hidden [[treasure]]. These spots are protected by a spell that prevents anyone but the fern-seed holder from ever knowing their locations.<ref>{{cite web |url=http://www.saunalahti.fi/~marian1/gourmet/season5a.htm |title=Traditional Finnish Midsummer celebration |publisher=Saunalahti.fi |access-date=2013-09-07 |archive-date=19 September 2011 |archive-url=https://web.archive.org/web/20110919201014/http://www.saunalahti.fi/~marian1/gourmet/season5a.htm }}</ref> In Wicca, ferns are thought to have magical properties such as a dried fern can be thrown into hot coals of a fire to exorcise evil spirits, or smoke from a burning fern is thought to drive away snakes and such creatures.<ref name="Cunningham">{{cite book |last=Cunningham |first=Scott |author-link=Scott Cunningham |title=Cunningham's Encyclopedia of Magical Herbs |date=1999 |publisher=Llewellyn |page=102}}</ref>
 === New Zealand ===
-Ferns are the national emblem of New Zealand and feature on its passport and in the design of its national airline, [[Air New Zealand]], and of its rugby team, the [[All Blacks]].
+The [[Silver fern]] is the national emblem of New Zealand and feature on its passport and in the design of its national airline, [[Air New Zealand]], and of its rugby team, the [[All Blacks]].{{cn|date=July 2025}}
 ==Organisms confused with ferns==
@@ Line 350: / Line 347: @@
 Several non-fern plants (and even animals) are called ferns and are sometimes confused with ferns. These include:
 * Asparagus fern—This may apply to one of several species of the [[monocotyledon|monocot]] genus ''[[Asparagus (genus)|Asparagus]]'', which are flowering plants.
 * Sweetfern—A flowering shrub of the genus ''[[Comptonia (plant)|Comptonia]]''.
@@ Line 382: / Line 378: @@
 == Bibliography ==
 {{Refbegin|30em}}
 === Books ===
+* {{cite book |last1=Christenhusz |first1=Maarten M. J. |last2=Fay |first2=Michael |last3=Byng |first3=James W. |author-link1=Maarten Christenhusz |author-link2=Michael Francis Fay |author-link3=James W. Byng |title=The Global Flora: Special Edition: GLOVAP Nomenclature Part 1 |url=https://books.google.com/books?id=DX60DwAAQBAJ |date=2018 |publisher=Plant Gateway Limited |isbn=978-0-9929993-6-0 |ref={{harvid|Christenhusz et al|2018}} }}
-* {{cite book |last1=Christenhusz |first1=Maarten M. J. |last2=Fay |first2=Michael |last3=Byng |first3=James W. |author-link1=Maarten Christenhusz |author-link2=Michael Francis Fay |author-link3=James W. Byng |title=The Global Flora: Special Edition: GLOVAP Nomenclature Part 1 |url=https://books.google.com/books?id=DX60DwAAQBAJ |date=2018 |publisher=Plant Gateway Limited |isbn=978-0-9929993-6-0 |ref={{harvid|Christenhusz et al|2018}}}}
+* {{cite book |last=Linnaeus |first=Carl |author-link=Carl Linnaeus |title=Species Plantarum: exhibentes plantas rite cognitas, ad genera relatas, cum differentiis specificis, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas |url=https://www.biodiversitylibrary.org/bibliography/669#/summary |year=1753 |volume=1 |chapter=Cryptogamia: Filices Musci |chapter-url=https://www.biodiversitylibrary.org/item/13830#page/503/mode/1up |pages=1061–1100, 1100–1130 |publisher=Impensis Laurentii Salvii |location=Stockholm, Sweden }}, ''see also'' [[Species Plantarum]]
-* {{cite book |last=Linnaeus |first=Carl |author-link=Carl Linnaeus |title=Species Plantarum: exhibentes plantas rite cognitas, ad genera relatas, cum differentiis specificis, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas |url=https://www.biodiversitylibrary.org/bibliography/669#/summary |year=1753 |volume=1 |chapter=Cryptogamia: Filices Musci |chapter-url=https://www.biodiversitylibrary.org/item/13830#page/503/mode/1up |pages=1061–1100, 1100–1130 |publisher=Impensis Laurentii Salvii |location=Stockholm, Sweden}}, ''see also'' [[Species Plantarum]]
 * Lord, Thomas R. (2006). ''Ferns and Fern Allies of Pennsylvania''. Indiana, Pennsylvania: Pinelands Press. [http://www.paferns.com/ Ferns and Fern Allies of Pennsylvania – Thomas Reeves Lord].
 * Moran, Robbin C. (2004). ''A Natural History of Ferns''. Portland, Oregon: Timber Press. {{ISBN|0-88192-667-1}}
-* {{cite book |last1=Ranker |first1=Tom A. |last2=Haufler |first2=Christopher H. |title=Biology and Evolution of Ferns and Lycophytes |url=https://books.google.com/books?id=js9JnwEACAAJ |date=2008 |publisher=[[Cambridge University Press]] |isbn=978-0-521-87411-3}}
+* {{cite book |last1=Ranker |first1=Tom A. |last2=Haufler |first2=Christopher H. |title=Biology and Evolution of Ferns and Lycophytes |url=https://books.google.com/books?id=js9JnwEACAAJ |date=2008 |publisher=[[Cambridge University Press]] |isbn=978-0-521-87411-3 }}
-* {{cite book |last=Swartz |first=Olof |author-link=Olof Swartz |title=Synopsis filicum: earum genera et species systematice complectens: adjectis lycopodineis, et descriptionibus novarum et rariorum specierum: cum tabulis aeneis quinque |date=1806 |publisher=Impensis Bibliopolii novi academici |location=[[Kiliae]] |url=https://www.biodiversitylibrary.org/bibliography/81948#/summary}}
+* {{cite book |last=Swartz |first=Olof |author-link=Olof Swartz |title=Synopsis filicum: earum genera et species systematice complectens: adjectis lycopodineis, et descriptionibus novarum et rariorum specierum: cum tabulis aeneis quinque |date=1806 |publisher=Impensis Bibliopolii novi academici |location=[[Kiliae]] |url=https://www.biodiversitylibrary.org/bibliography/81948#/summary }}
 === Journal articles ===
+* {{cite journal |last=Berry |first=Chris |title=The Middle Devonian plant collections of Francois Stockmans reconsidered. |journal=[[Geologica Belgica]] |date=2009 |volume=12 |issue=1–2 |pages=25–30 |url=https://popups.uliege.be/1374-8505/index.php?id=2633&file=1 }}
-* {{cite journal |last=Berry |first=Chris |title=The Middle Devonian plant collections of Francois Stockmans reconsidered. |journal=[[Geologica Belgica]] |date=2009 |volume=12 |issue=1–2 |pages=25–30 |url=https://popups.uliege.be/1374-8505/index.php?id=2633&file=1}}
+* {{cite journal |last1=Bomfleur |first1=B. |last2=McLoughlin |first2=S. |last3=Vajda |first3=V. |title=Fossilized Nuclei and Chromosomes Reveal 180 Million Years of Genomic Stasis in Royal Ferns |url=http://nrm.diva-portal.org/smash/get/diva2:719316/FULLTEXT01 |journal=[[Science (journal)|Science]] |date=20 March 2014 |volume=343 |issue=6177 |pages=1376–1377 |doi=10.1126/science.1249884 |pmid=24653037 |bibcode=2014Sci...343.1376B |s2cid=38248823 |ref={{harvid|Bomfleur et al|2014}} }}
-* {{cite journal |last1=Bomfleur |first1=B. |last2=McLoughlin |first2=S. |last3=Vajda |first3=V. |title=Fossilized Nuclei and Chromosomes Reveal 180 Million Years of Genomic Stasis in Royal Ferns |url=http://nrm.diva-portal.org/smash/get/diva2:719316/FULLTEXT01 |journal=[[Science (journal)|Science]] |date=20 March 2014 |volume=343 |issue=6177 |pages=1376–1377 |doi=10.1126/science.1249884 |pmid=24653037 |bibcode=2014Sci...343.1376B |s2cid=38248823 |ref={{harvid|Bomfleur et al|2014}}}}
+* {{cite journal |last1=Cantino |first1=Philip D. |last2=Doyle |first2=James A. |last3=Graham |first3=Sean W. |last4=Judd |first4=Walter S. |author-link4=Walter S. Judd |last5=Olmstead |first5=Richard G. |last6=Soltis |first6=Douglas E. |author-link6=Douglas Soltis |last7=Soltis |first7=Pamela S. |author-link7=Pamela Soltis |last8=Donoghue |first8=Michael J. |author-link8=Michael Donoghue |title=Towards a Phylogenetic Nomenclature of Tracheophyta |journal=[[Taxon (journal)|Taxon]] |date=1 August 2007 |volume=56 |issue=3 |page=822 |doi=10.2307/25065865 |ref={{harvid|Cantino et al|2007}} |jstor=25065865}}
-* {{cite journal |last1=Cantino |first1=Philip D. |last2=Doyle |first2=James A. |last3=Graham |first3=Sean W. |last4=Judd |first4=Walter S. |author-link4=Walter S. Judd |last5=Olmstead |first5=Richard G. |last6=Soltis |first6=Douglas E. |author-link6=Douglas Soltis |last7=Soltis |first7=Pamela S. |author-link7=Pamela Soltis |last8=Donoghue |first8=Michael J. |author-link8=Michael Donoghue |title=Towards a Phylogenetic Nomenclature of Tracheophyta |journal=[[Taxon (journal)|Taxon]] |date=1 August 2007 |volume=56 |issue=3 |pages=822 |doi=10.2307/25065865 |ref={{harvid|Cantino et al|2007}} |jstor=25065865}}
 * {{cite journal |last1=Chase |first1=Mark W. |author-link1=Mark W. Chase |last2=Reveal |first2=James L. |author-link2=James L. Reveal |name-list-style=amp |year=2009 |title=A phylogenetic classification of the land plants to accompany APG III |journal=[[Botanical Journal of the Linnean Society]] |volume=161 |issue=2 |pages=122–127 |doi=10.1111/j.1095-8339.2009.01002.x |doi-access=free}}
-* {{cite journal |last1=Christenhusz |first1=Maarten J. M. |last2=Byng |first2=J. W. |author-link1=Maarten Christenhusz |author-link2=James W. Byng |name-list-style=amp |year=2016 |title=The number of known plants species in the world and its annual increase |journal=[[Phytotaxa]] |volume=261 |pages=201–217 |url=http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 |doi=10.11646/phytotaxa.261.3.1 |issue=3 |publisher=Magnolia Press |doi-access=free}}
+* {{cite journal |last1=Christenhusz |first1=Maarten J. M. |last2=Byng |first2=J. W. |author-link1=Maarten Christenhusz |author-link2=James W. Byng |name-list-style=amp |year=2016 |title=The number of known plants species in the world and its annual increase |journal=[[Phytotaxa]] |volume=261 |pages=201–217 |url=http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 |doi=10.11646/phytotaxa.261.3.1 |issue=3 |publisher=Magnolia Press |doi-access=free |bibcode=2016Phytx.261..201C }}
-* {{cite journal |last1=Christenhusz |first1=Maarten J. M. |author-link1=Maarten J. M. Christenhusz |last2=Zhang |first2=X. C. |last3=Schneider |first3=H. |title=A linear sequence of extant families and genera of lycophytes and ferns |journal=[[Phytotaxa]] |date=18 February 2011 |volume=19 |issue=1 |pages=7 |doi=10.11646/phytotaxa.19.1.2 |ref={{harvid|Christenhusz et al|2011}} |doi-access=free|hdl=10138/28042 |hdl-access=free }}
+* {{cite journal |last1=Christenhusz |first1=Maarten J. M. |author-link1=Maarten J. M. Christenhusz |last2=Zhang |first2=X. C. |last3=Schneider |first3=H. |title=A linear sequence of extant families and genera of lycophytes and ferns |journal=[[Phytotaxa]] |date=18 February 2011 |volume=19 |issue=1 |page=7 |doi=10.11646/phytotaxa.19.1.2 |ref={{harvid|Christenhusz et al|2011}} |doi-access=free|bibcode=2011Phytx..19....7C |hdl=10138/28042 |hdl-access=free }}
 * {{cite journal |last1=Christenhusz |first1=Maarten J. M. |author-link1=Christenhusz |last2=Chase |first2=Mark W. |author-link2=Mark W. Chase |year=2014 |title=Trends and concepts in fern classification |journal=[[Annals of Botany]] |volume=113 |issue=4 |pages=571–594 |doi=10.1093/aob/mct299 |pmid=24532607 |pmc=3936591}}
-* {{cite journal |last1=Christenhusz |first1=Maarten J. M. |last2=Chase |first2=Mark W. |author-link1=Maarten Christenhusz |author-link2=Mark W. Chase |title=PPG recognises too many fern genera |journal=[[Taxon (journal)|Taxon]] |date=1 June 2018 |volume=67 |issue=3 |pages=481–487 |doi=10.12705/673.2 |doi-access=free}}
+* {{cite journal |last1=Christenhusz |first1=Maarten J. M. |last2=Chase |first2=Mark W. |author-link1=Maarten Christenhusz |author-link2=Mark W. Chase |title=PPG recognises too many fern genera |journal=[[Taxon (journal)|Taxon]] |date=1 June 2018 |volume=67 |issue=3 |pages=481–487 |doi=10.12705/673.2 |doi-access=free|bibcode=2018Taxon..67..481C }}
 * {{Cite journal |last1=May |first1=Lenore Wile |title=The economic uses and associated folklore of ferns and fern allies |journal=[[The Botanical Review]] |year=1978 |volume=44 |issue=4 |pages=491–528 |doi=10.1007/BF02860848 |bibcode=1978BotRv..44..491M |s2cid=42101599}}
 * {{cite journal |last1=Melan |first1=M. A. |last2=Whittier |first2=D. P. |year=1990 |title=Effects of Inorganic Nitrogen Sources on Spore Germination and Gametophyte Growth in Botrychium Dissectum |journal=Plant, Cell and Environment |volume=13 |issue=5 |pages=477–482 |doi=10.1111/j.1365-3040.1990.tb01325.x |bibcode=1990PCEnv..13..477M}}
 * {{cite journal |last1=Pryer |first1=Kathleen M. |last2=Schneider |first2=Harald |last3=Smith |first3=Alan R. |last4=Cranfill |first4=Raymond |last5=Wolf |first5=Paul G. |last6=Hunt |first6=Jeffrey S. |last7=Sipes |first7=Sedonia D. |year=2001 |title=Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants |journal=Nature |volume=409 |issue=6820 |pages=618–622 |doi=10.1038/35054555 |pmid=11214320 |bibcode=2001Natur.409..618S |s2cid=4367248}}
-* {{cite journal |last1=Pryer |first1=Kathleen M. |last2=Schuettpelz |first2=Eric |last3=Wolf |first3=Paul G. |last4=Schneider |first4=Harald |last5=Smith |first5=Alan R. |last6=Cranfill |first6=Raymond |year=2004 |title=Phylogeny and evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences |journal=American Journal of Botany |volume=91 |issue=10 |pages=1582–1598 |doi=10.3732/ajb.91.10.1582 |pmid=21652310}}
+* {{cite journal |last1=Pryer |first1=Kathleen M. |last2=Schuettpelz |first2=Eric |last3=Wolf |first3=Paul G. |last4=Schneider |first4=Harald |last5=Smith |first5=Alan R. |last6=Cranfill |first6=Raymond |year=2004 |title=Phylogeny and evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences |journal=American Journal of Botany |volume=91 |issue=10 |pages=1582–1598 |doi=10.3732/ajb.91.10.1582 |pmid=21652310|bibcode=2004AmJB...91.1582P }}
-* {{cite journal |last1=Pteridophyte Phylogeny Group |title=A community-derived classification for extant lycophytes and ferns |journal=Journal of Systematics and Evolution |date=November 2016 |volume=54 |issue=6 |pages=563–603 |doi=10.1111/jse.12229 |s2cid=39980610 |doi-access=free}}
+* {{cite journal |last1=Pteridophyte Phylogeny Group |title=A community-derived classification for extant lycophytes and ferns |journal=Journal of Systematics and Evolution |date=November 2016 |volume=54 |issue=6 |pages=563–603 |doi=10.1111/jse.12229 |s2cid=39980610 |doi-access=free|bibcode=2016JSyEv..54..563. }}
-* {{cite journal |last1=Schneider |first1=Harald |last2=Smith |first2=Alan R. |last3=Pryer |first3=Kathleen M. |title=Is Morphology Really at Odds with Molecules in Estimating Fern Phylogeny? |journal=[[Systematic Botany]] |date=1 July 2009 |volume=34 |issue=3 |pages=455–475 |doi=10.1600/036364409789271209 |s2cid=85855934 |ref={{harvid|Schneider et al|2009}}}}
+* {{cite journal |last1=Schneider |first1=Harald |last2=Smith |first2=Alan R. |last3=Pryer |first3=Kathleen M. |title=Is Morphology Really at Odds with Molecules in Estimating Fern Phylogeny? |journal=[[Systematic Botany]] |date=1 July 2009 |volume=34 |issue=3 |pages=455–475 |doi=10.1600/036364409789271209 |bibcode=2009SysBo..34..455S |s2cid=85855934 |ref={{harvid|Schneider et al|2009}}}}
-* {{Cite thesis |last=Schuettpelz |first=Eric |title=The evolution and diversification of epiphytic ferns |url=http://dukespace.lib.duke.edu/dspace/bitstream/10161/181/1/D_Schuettpelz_Eric_a_052007.pdf |year=2007 |publisher=[[Duke University]] |type=[[PhD thesis]] |contribution=Table 1 |access-date=11 December 2009 |archive-date=20 June 2010 |archive-url=https://web.archive.org/web/20100620022844/http://dukespace.lib.duke.edu/dspace/bitstream/10161/181/1/D_Schuettpelz_Eric_a_052007.pdf |url-status=dead}}
+* {{Cite thesis |last=Schuettpelz |first=Eric |title=The evolution and diversification of epiphytic ferns |url=http://dukespace.lib.duke.edu/dspace/bitstream/10161/181/1/D_Schuettpelz_Eric_a_052007.pdf |year=2007 |publisher=[[Duke University]] |type=[[PhD thesis]] |contribution=Table 1 |access-date=11 December 2009 |archive-date=20 June 2010 |archive-url=https://web.archive.org/web/20100620022844/http://dukespace.lib.duke.edu/dspace/bitstream/10161/181/1/D_Schuettpelz_Eric_a_052007.pdf }}
-* {{cite journal |last1=Schuettpelz |first1=Eric |last2=Rouhan |first2=Germinal |last3=Pryer |first3=Kathleen M. |last4=Rothfels |first4=Carl J. |last5=Prado |first5=Jefferson |last6=Sundue |first6=Michael A. |last7=Windham |first7=Michael D. |last8=Moran |first8=Robbin C. |last9=Smith |first9=Alan R. |title=Are there too many fern genera? |journal=[[Taxon (journal)|Taxon]] |date=1 June 2018 |volume=67 |issue=3 |pages=473–480 |doi=10.12705/673.1 |ref={{harvid|Schuettpelz et al|2018}} |doi-access=free}}
+* {{cite journal |last1=Schuettpelz |first1=Eric |last2=Rouhan |first2=Germinal |last3=Pryer |first3=Kathleen M. |last4=Rothfels |first4=Carl J. |last5=Prado |first5=Jefferson |last6=Sundue |first6=Michael A. |last7=Windham |first7=Michael D. |last8=Moran |first8=Robbin C. |last9=Smith |first9=Alan R. |title=Are there too many fern genera? |journal=[[Taxon (journal)|Taxon]] |date=1 June 2018 |volume=67 |issue=3 |pages=473–480 |doi=10.12705/673.1 |ref={{harvid|Schuettpelz et al|2018}} |doi-access=free|bibcode=2018Taxon..67..473S }}
-* {{cite journal |last=Smith |first=Alan R. |author2=Kathleen M. Pryer |author3=Eric Schuettpelz |author4=Petra Korall |author5=Harald Schneider |author6=Paul G. Wolf |year=2006 |title=A classification for extant ferns |url=https://www.idigbio.org/wiki/images/9/95/Smith_et_al_2006.pdf |journal=[[Taxon (journal)|Taxon]] |volume=55 |issue=3 |pages=705–731 |doi=10.2307/25065646 |jstor=25065646 |ref={{harvid|Smith et al.2006}}}}
+* {{cite journal |last=Smith |first=Alan R. |author2=Kathleen M. Pryer |author3=Eric Schuettpelz |author4=Petra Korall |author5=Harald Schneider |author6=Paul G. Wolf |year=2006 |title=A classification for extant ferns |url=https://www.idigbio.org/wiki/images/9/95/Smith_et_al_2006.pdf |journal=[[Taxon (journal)|Taxon]] |volume=55 |issue=3 |pages=705–731 |doi=10.2307/25065646 |jstor=25065646 |bibcode=2006Taxon..55..705S |ref={{harvid|Smith et al.2006}} }}
 * {{cite journal |last1=Stein |first1=W. E. |last2=Mannolini |first2=F. |last3=Hernick |first3=L. V. |last4=Landling |first4=E. |last5=Berry |first5=C. M. |year=2007 |title=Giant cladoxylopsid trees resolve the enigma of the Earth's earliest forest stumps at Gilboa |journal=[[Nature (journal)|Nature]] |volume=446 |issue=7138 |pages=904–907 |doi=10.1038/nature05705 |pmid=17443185 |bibcode=2007Natur.446..904S |s2cid=2575688 |ref={{harvid|Stein et al|2007}}}}
-* {{cite journal |author=Walkowiak |first=Radoslaw Janusz |title=Classification of Pteridophytes – Short classification of the ferns. |journal=IEA Paper |date=2017 |url=https://internationalequisetologicalassociation.yolasite.com/resources/Classification%20of%20Pteridophytes.pdf |doi=10.13140/RG.2.2.29934.20809}}
+* {{cite journal |author=Walkowiak |first=Radoslaw Janusz |title=Classification of Pteridophytes – Short classification of the ferns. |journal=IEA Paper |date=2017 |url=https://internationalequisetologicalassociation.yolasite.com/resources/Classification%20of%20Pteridophytes.pdf |doi=10.13140/RG.2.2.29934.20809 }}
 * {{cite journal |last1=Underwood |first1=L. M. |title=The early writers on ferns and their collections. I. Linnaeus, 1707–1778 |journal=[[Torrey Botanical Society|Torreya]] |date=1903 |volume=3 |issue=10 |pages=145–150 |jstor=40594126 |issn=0096-3844}}
 === Websites ===
+* {{cite web |last1=McCausland |first1=Jim |title=Rediscover ferns |url=https://www.sunset.com/garden/flowers-plants/rediscover-ferns |website=Garden plants |publisher=Sunset Magazine |access-date=22 November 2019 |date=22 February 2019 |archive-date=30 October 2021 |archive-url=https://web.archive.org/web/20211030104351/https://www.sunset.com/garden/flowers-plants/rediscover-ferns }}
-* {{cite web |last1=McCausland |first1=Jim |title=Rediscover ferns |url=https://www.sunset.com/garden/flowers-plants/rediscover-ferns |website=Garden plants |publisher=Sunset Magazine |access-date=22 November 2019 |date=22 February 2019 |archive-date=30 October 2021 |archive-url=https://web.archive.org/web/20211030104351/https://www.sunset.com/garden/flowers-plants/rediscover-ferns |url-status=dead}}
+* {{cite web |title=Pteridopsida: Fossil Record |url=http://www.ucmp.berkeley.edu/plants/pterophyta/pteridofr.html |website=Plants: Pteridopsida |publisher=[[University of California Museum of Paleontology]] |access-date=23 November 2019 |ref={{harvid|UCMP|2019}} }}
-* {{cite web |title=Pteridopsida: Fossil Record |url=http://www.ucmp.berkeley.edu/plants/pterophyta/pteridofr.html |website=Plants: Pteridopsida |publisher=[[University of California Museum of Paleontology]] |access-date=23 November 2019 |ref={{harvid|UCMP|2019}}}}
+* {{cite web |title=Classifying and identifying ferns |url=https://www.sciencelearn.org.nz/resources/1104-classifying-and-identifying-ferns |website=Science Learning Hub |publisher=[[The University of Waikato]] |access-date=24 November 2019 |language=en |date=3 September 2018 |ref={{harvid|SLH|2018}} }}
-* {{cite web |title=Classifying and identifying ferns |url=https://www.sciencelearn.org.nz/resources/1104-classifying-and-identifying-ferns |website=Science Learning Hub |publisher=[[The University of Waikato]] |access-date=24 November 2019 |language=en |date=3 September 2018 |ref={{harvid|SLH|2018}}}}
+* {{cite encyclopedia |last1=Mickel |first1=John T. |last2=Wagner |first2=Warren H. |last3=Gifford |first3=Ernest M. |display-authors=etal |title=Fern |url=https://www.britannica.com/plant/fern |encyclopedia=[[Encyclopædia Britannica]] |access-date=24 November 2019 |date=4 February 2019 |ref={{harvid|EB|2019}} }}
-* {{cite encyclopedia |last1=Mickel |first1=John T. |last2=Wagner |first2=Warren H. |last3=Gifford |first3=Ernest M. |display-authors=etal |title=Fern |url=https://www.britannica.com/plant/fern |encyclopedia=[[Encyclopædia Britannica]] |access-date=24 November 2019 |date=4 February 2019 |ref={{harvid|EB|2019}}}}
+* {{cite web |last1=Hassler |first1=Michael |last2=Schmitt |first2=Bernd |title=Checklist of Ferns and Lycophytes of the World |url=https://worldplants.webarchiv.kit.edu/ferns/ |website=World Ferns |publisher=[[Botanical Garden of the Karlsruhe Institute of Technology]] |access-date=25 November 2019 |date=2 November 2019 |archive-date=2 September 2017 |archive-url=https://web.archive.org/web/20170902225743/http://worldplants.webarchiv.kit.edu/ferns/ }}
-* {{cite web |last1=Hassler |first1=Michael |last2=Schmitt |first2=Bernd |title=Checklist of Ferns and Lycophytes of the World |url=https://worldplants.webarchiv.kit.edu/ferns/ |website=World Ferns |publisher=[[Botanical Garden of the Karlsruhe Institute of Technology]] |access-date=25 November 2019 |date=2 November 2019 |archive-date=2 September 2017 |archive-url=https://web.archive.org/web/20170902225743/http://worldplants.webarchiv.kit.edu/ferns/ |url-status=dead}}
+* {{cite web |last1=Pryer |first1=Kathleen M. |last2=Smith |first2=Alan R. |last3=Rothfels |first3=Carl |title=Polypodiopsida |url=http://tolweb.org/Polypodiopsida/20615 |website=[[Tree of Life]] |date=2009 |ref={{harvid|Pryer et al|2006}} }}
-* {{cite web |last1=Pryer |first1=Kathleen M. |last2=Smith |first2=Alan R. |last3=Rothfels |first3=Carl |title=Polypodiopsida |url=http://tolweb.org/Polypodiopsida/20615 |website=[[Tree of Life]] |date=2009 |ref={{harvid|Pryer et al|2006}}}}
 * [https://web.archive.org/web/20160304060354/http://www.anbg.gov.au/fern/taxa/classification.html ''A classification of the ferns and their allies''.] (Australian National Herbarium)
 * [https://web.archive.org/web/20160117055319/http://www.jaknouse.athens.oh.us/ferns/bookfern.html ''A fern book bibliography''.]
@@ Line 430: / Line 422: @@
 * [https://web.archive.org/web/20120512162340/http://www.cas.vanderbilt.edu/bioimages/pages/non-seed-plants.htm Non-seed plant images at ''bioimages.vanderbilt.edu'']
 * [http://www.amerfernsoc.org/ American Fern Society]
-* [http://www.eBPS.org.uk/ British Pteridological Society]
+* [http://www.eBPS.org.uk/ British Pteridological Society] {{Webarchive|url=https://web.archive.org/web/20190102094708/https://ebps.org.uk/ |date=2 January 2019 }}
 * [http://internationalequisetologicalassociation.yolasite.com/ International Equisetological Association]
 {{Refend}}

Fern: Difference between revisions

Latest revision as of 04:42, 20 November 2025

Description

Sporophyte

Gametophyte

Life cycle and reproduction

Taxonomy

Molecular phylogenetics

Phylogeny

Nomenclature and subdivision

Evolution and biogeography

Distribution and habitat

Ecology

Life cycle

Uses

Culture

Pteridology

Pteridomania

Other applications

Folklore

New Zealand

Organisms confused with ferns

Misnomers

Fern-like flowering plants

See also

Notes

References

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