Fern

Sporophyte Extant ferns are herbaceous perennials and most lack woody growth. When woody growth is present, it is found in the stem. Their foliage may be deciduous or evergreen, and some are semi-evergreen depending on the climate. Like the sporophytes of seed plants, those of ferns consist of stems, leaves and roots. Ferns differ from spermatophytes in that they reproduce by spores rather than by seeds. 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. The fertile leaves are much narrower than the sterile leaves, and may have no green tissue at all, as in the Blechnaceae and Lomariopsidaceae. 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 (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. 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. 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 tall in a few species (e.g., Cyathea brownii on Norfolk Island and Cyathea medullaris in New Zealand). Roots are underground non-photosynthetic structures that take up water and nutrients from soil. They are always fibrous and are structurally very similar to the roots of seed plants. 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. A fern gametophyte typically consists of: usually heart or kidney shaped, 3–10 mm long and 2–8 mm broad. The prothallus produces gametes by means of: • Antheridia: Small spherical structures that produce flagellate antherozoids. • Archegonia: A flask-shaped structure that produces a single egg at the bottom, reached by the male gametophyte by swimming down the neck. greatly elongated cells, that absorb water and mineral salts over the whole structure. Rhizoids anchor the prothallus to the soil. == Life cycle and reproduction ==

'') 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 ( 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. 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. 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. of monarch fern contain clusters of sporangia == 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). By 1806 this had increased to 38 genera, and has progressively increased since (see ). 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. 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: • 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. The leptosporangiate ferns are sometimes called "true ferns". This group includes most plants familiarly known as ferns. Modern research supports older ideas based on morphology that the Osmundaceae diverged early in the evolutionary history of the leptosporangiate ferns; in certain ways this family is intermediate between the eusporangiate ferns and the leptosporangiate ferns. Rai and Graham (2010) broadly supported the primary groups, but queried their relationships, concluding that "at present perhaps the best that can be said about all relationships among the major lineages of monilophytes in current studies is that we do not understand them very well". Grewe et al. (2013) confirmed the inclusion of horsetails within ferns sensu lato, but also suggested that uncertainties remained in their precise placement. The phylogenetic relationships are shown in the following cladogram (to the level of orders). 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. 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. == 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. New Zealand, for which the fern is a symbol, has about 230 species, distributed throughout the country. 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). The epiphytic ferns in particular 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. 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 usually found 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. File:CowellFerns.JPG|In undergrowth below coast redwoods, California File:FernBedInForest.jpg|Fern bed under a forest canopy, Virginia File:Samambaia fern.jpg|On a wall File:Harts Tongue Fern on the Great Orme - geograph.org.uk - 345785.jpg|hart's tongue fern Asplenium scolopendrium in a gryke in limestone pavement File:... epiphytic fern (49095803342).jpg|Epiphytic ferns in India File:Canning_rv_azolla_10_gnangarra.jpg|Azolla duckweed fern coveringthe Canning River, Western Australia == 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) 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. Fern tubers were used by the Guanches to make gofio in the Canary Islands. Ferns are generally not known to be poisonous to humans. Licorice fern rhizomes were chewed by the natives of the Pacific Northwest for their flavor. 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. 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. 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. Several ferns, such as bracken and Azolla 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. The important fossil fuel coal consists of the remains of primitive plants, including ferns. == Culture ==

: 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. Other applications created using a chaos game, through an Iterated function system 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. 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. 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. 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. ==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 flowering plants in the monocot genus Asparagus. • 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 and gymnosperms 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. Cycads, which are gymnosperms also have pinnate leaves that somewhat resemble fern fronds. == See also ==