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The ferns ( Polypodiopsida or Polypodiophyta) are a group of (plants with xylem and phloem) that reproduce via and have neither nor . They differ from 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. (2025). 9780143419549, DK Pub.. ISBN 9780143419549
Ferns have complex leaf called that are more complex than the of clubmosses. Most ferns are leptosporangiate ferns. They produce coiled Fiddlehead fern that uncoil and expand into . 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 , Psilotaceae, , and ophioglossoid ferns.
The fern crown group, consisting of the leptosporangiates and eusporangiates, is estimated to have originated in the late Silurian period 423.2 million years ago, but Polypodiales, the group that makes up 80% of living fern diversity, did not appear and diversify until 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 Paddy field. They also play certain roles in folklore.
The green, photosynthesis part of the plant is technically a 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 into . This uncurling of the leaf is termed circinate vernation. Leaves are divided into two types: sporophylls and tropophylls. produce spores; do not. Fern spores are borne in sporangia which are usually clustered to form sorus. The sporangia may be covered with a protective coating called an indusium. The arrangement of the sporangia is important in classification.
In monomorphic ferns, the fertile and sterile leaves look morphologically the same, and both are able to photosynthesize. In hemidimorphic ferns, just a portion of the fertile leaf is different from the sterile leaves. In dimorphic (holomorphic) ferns, the two types of leaves are frond dimorphism. Understanding the contribution of LFY and PEBP flowering genes to fern leaf dimorphism – Botany 2019. The fertile leaves are much narrower than the sterile leaves, and may have no green tissue at all, as in the Blechnaceae and Lomariopsidaceae.
The anatomy of fern leaves can be anywhere from simple to highly divided, or even indeterminate (e.g. Gleicheniaceae, Lygodium). The divided forms are Pinnation, 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. Fern Structure – Forest Service. Fern Structure – Forest Service, Auckland, New Zealand. 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 , even though they grow underground only in some of the species. Epiphytic species and many of the terrestrial ones have above-ground creeping (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.
Traditionally, three discrete groups have been denominated ferns: two groups of eusporangiate ferns, the families Ophioglossaceae (Ophioglossum, , 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 sister group to the leptosporangiate ferns. Several other groups of species were considered fern allies: the , , and quillworts in Lycopodiophyta; the whisk ferns of Psilotaceae; and the horsetails of Equisetaceae. Since this grouping is Polyphyly, 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 , 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 and Marattiaceae are arguably another clade.
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"., p. 1450 Grewe et al. (2013) confirmed the inclusion of horsetails within ferns sensu lato, but also suggested that uncertainties remained in their precise placement. Other classifications have raised Ophioglossales to the rank of a fifth class, separating the whisk ferns and ophioglossoid ferns.
In addition they defined 11 orders and 37 families. That system was a consensus of a number of studies, and was further refined. The phylogenetic relationships are shown in the following cladogram (to the level of orders). This division into four major clades was then confirmed using plant morphology alone.
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). Christenhusz et al. (2011) followed this use of subclasses but recombined Smith's Psilotopsida as Ophioglossidae, giving four subclasses of ferns again.
Christenhusz and Mark 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;
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 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:
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. 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.) 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.
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 , a bryophyte lineage.
Due to the very large genome seen in most ferns, it was suspected they might have gone through Paleopolyploidy, 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. Genes for seeds arose early in plant evolution, ferns reveal
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.
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.
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 , 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.
The life cycle of a typical fern proceeds as follows:
Sometimes a gametophyte can give rise to sporophyte traits like roots or sporangia without the rest of the sporophyte. The Ferns (Filicales): Volume 1, Analytical Examination of the Criteria of Comparison: Treated Comparatively with a View to their Natural Classification
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 foliage and as , especially the Boston fern ( Nephrolepis exaltata) and other members of the genus Nephrolepis. The Asplenium nidus ( Asplenium nidus) is also popular, as are the (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 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.
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.
were popular in America in the 1970s and 80s.
Phylogeny
The ferns are related to other groups as shown in the following cladogram:
Nomenclature and subdivision
The classification of Smith et al. in 2006 treated ferns as four classes:
+ Comparison of fern subdivisions in some classifications Class Polypodiopsida Subclass Equisetidae Subclass Ophioglossidae Subclass Marattiidae 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. Ferns evolved to cope with low-light conditions present under the canopy of angiosperms.
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 forests.
Ecology
Fern species live in a wide variety of , 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 . 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 ; crevices in rock faces, especially when sheltered from the full sun; acid wetlands including and ; and tropical , where many species are (something like a quarter to a third of all fern species).
the Canning River, Western Australia
Life cycle
Ferns are differing from by having true leaf (megaphylls), which are often pinnate. They differ from spermatophyte ( and ) in reproducing by means of spores and lacking and . Like all embryophyte, they have a life cycle referred to as alternation of generations, characterized by alternating diploid sporophyte and haploid gametophyte phases. The diploid sporophyte has 2 n paired , 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.
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 Oceania. Fern tubers were used for food 30,000 years ago in Europe.Van Gilder Cooke, Sonia (23 October 2010). "Stone Age humans liked their burgers in a bun". New Scientist, p. 18.Revedin, Anna et al. (18 October 2010). "Thirty thousand-year-old evidence of plant food processing". PNAS. 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 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.
Culture
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 .
Pteridomania
Pteridomania was a Victorian era craze which involved fern collecting and fern motifs in decorative art including pottery, glass, , , wood, printing, and sculpture "appearing on everything from infant baptism presents to 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
The Barnsley fern is a fractal named after the British mathematician Michael Barnsley who first described it in his book Fractals Everywhere. A Self-similarity structure is described by a mathematical function, applied repeatedly at different scales to create a frond pattern. Fractals Everywhere, Boston, MA: Academic Press, 1993,
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, Finland 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
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.
Organisms confused with ferns
Misnomers
Several non-fern plants (and even animals) are called ferns and are sometimes confused with ferns. These include:
Fern-like flowering plants
Some such as Arecaceae and members of the Apiaceae 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.
See also
Notes
Bibliography
Books
Journal articles
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