A fern is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissues that conduct water and nutrients, in having branched stems and in having life cycles in which the sporophyte is the dominant phase. Like other vascular plants, ferns have complex leaves called megaphylls, that are more complex than the microphylls of clubmosses. Most ferns are leptosporangiate ferns, sometimes referred to as true ferns. They produce coiled fiddleheads that uncoil and expand into fronds.[3] The group includes about 10,560 known extant species.[4]

Ferns are defined here in the broad sense, being all of the Polypodiopsida, comprising both the leptosporangiate (Polypodiidae) and eusporangiate ferns, the latter itself comprising ferns other than those denominated true ferns, including horsetails or scouring rushes, whisk ferns, marattioid ferns, and ophioglossoid ferns.

Ferns first appear in the fossil record about 360 million years ago in the late Devonian period,[5] but many of the current families and species did not appear until roughly 145 million years ago in the early Cretaceous, after flowering plants came to dominate many environments. The fern Osmunda claytoniana is a paramount example of evolutionary stasis; paleontological evidence indicates it has remained unchanged, even at the level of fossilized nuclei and chromosomes, for at least 180 million years.[6]

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 are significant weeds. They also play certain roles in mythology and art.

Temporal range: Late Devonian[1]Present
Athyrium filix-femina
A fern unrolling a young frond
Scientific classification
Kingdom: Plantae
Subkingdom: Embryophyta
(unranked): Tracheophyta
(unranked): euphyllophytes
Class: Polypodiopsida
  • Monilophyta
  • Polypodiophyta
  • Filicophyta
  • Filices


Ferns at melb botanical gardens
Ferns at the Royal Melbourne Botanical Gardens
Tree ferns, probably Dicksonia antarctica, growing in Nunniong, Australia

Like the sporophytes of seed plants, those of ferns consist of stems, leaves and roots.

Stems: Fern stems are often referred to as 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). These can reach up to 20 meters (66 ft) tall in a few species (e.g., Cyathea brownii on Norfolk Island and Cyathea medullaris in New Zealand).[7]

Leaf: The green, photosynthetic part of the plant is technically a megaphyll and in ferns, it is often referred to as a frond. New leaves typically expand by the unrolling of a tight spiral called a crozier or fiddlehead fern. This uncurling of the leaf is termed circinate vernation. Leaves are divided into two types a trophophyll and a sporophyll. A trophophyll frond is a vegetative leaf analogous to the typical green leaves of seed plants that does not produce spores, instead only producing sugars by photosynthesis. A sporophyll frond is a fertile leaf that produces spores borne in sporangia that are usually clustered to form sori. In most ferns, fertile leaves are morphologically very similar to the sterile ones, and they photosynthesize in the same way. In some groups, the fertile leaves are much narrower than the sterile leaves, and may even have no green tissue at all (e.g., Blechnaceae, Lomariopsidaceae). The anatomy of fern leaves can either be simple or highly divided. In tree ferns, the main stalk that connects the leaf to the stem (known as the stipe), often have multiple leaflets. The leafy structures that grow from the stipe are known as pinnae and are often again divided into smaller pinnules.[8]

Roots: The underground non-photosynthetic structures that take up water and nutrients from soil. They are always fibrous and structurally are very similar to the roots of seed plants.

Like all other vascular plants, the diploid 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:

    • Antheridia: Small spherical structures that produce flagellate sperm.
  • Rhizoids: 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.


Ferns first appear in the fossil record in the early Carboniferous period. 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 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.[9] 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 much true 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:[10]

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.[10] However, there remained differences of opinion about the placement of the genus Equisetum (see Equisetopsida for further discussion). One possible solution was to only denominate the leptosporangiate ferns as true ferns while denominating the other three groups as fern allies. In practice, numerous classification schemes have been proposed for ferns and fern allies, and there has been little consensus among them.

The leptosporangiate ferns are sometimes called true ferns.[11] 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".[12] Grewe et al. (2013) confirmed the inclusion of horsetails within ferns sensu lato, but also suggested that uncertainties remained in their precise placement.[13] Other classifications have raised Ophioglossales to the rank of a fifth class, separating the whisk ferns and ophioglossoid ferns.[13]

One problem with the classification of ferns is that of cryptic species. A cryptic species is a species that is morphologically similar to another species, but differs genetically in ways that prevent fertile interbreeding. A good example of this is the currently designated species Asplenium trichomanes (maidenhair spleenwort). This is actually a species complex that includes distinct diploid and tetraploid races. There are minor but unclear morphological differences between the two groups, which prefer distinctly differing habitats. In many cases such as this, the species complexes have been separated into separate species, thus raising the total number of species of ferns. Possibly many more cryptic species are yet to be discovered and designated.


The ferns are related to other higher order taxa, as shown in the following cladogram:[9][14][15][2]

Tracheophyta - vascular plants

Lycopodiophyta (Lycopodiopsida) - lycophytes


Polypodiophyta (Polypodiopsida) - ferns

Spermatophyta - seed plants


Angiospermae - flowering plants


Smith's 2006 classification treated the ferns as four classes:[16][17]

In addition they defined 11 orders and 37 families.[16] That system was a consensus of a number of studies, and was further refined.[13][18] The phylogenetic relationships are shown in the following cladogram (to the level of orders).[19] This division into four major clades was then confirmed using morphology alone.[20]


Lycopodiophytes (club mosses, spike mosses, quillworts)


Spermatophytes (seed plants)


Psilotales (whisk ferns)  

Ophioglossales (grapeferns etc.)  


Equisetales (horsetails)  





Hymenophyllales (filmy ferns)  



Salviniales (heterosporous)  

Cyatheales (tree ferns)  


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).[15] Christenhusz et al.[a] (2011) followed this use of subclasses but recombined Smith's Psilotopsida as Ophioglossidae, giving four subclasses of ferns again.[21]

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;[9]

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:[2]






In the Pteridophyte Phylogeny Group classification 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) needs to be distinguished from the narrower usage (sensu stricto) of Smith et al. (Polypodiopsida sensu Smith et al.)[2]


Ferns at Muir Woods, California

The stereotypical image of ferns growing in moist shady woodland nooks is far from a complete picture of the habitats where ferns can be found growing. Fern species live in a wide variety of habitats, from remote mountain elevations, to dry desert rock faces, to bodies of water or in 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[22]).

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.[23]

Many ferns depend on associations with mycorrhizal fungi. Many ferns only grow within specific pH ranges; for instance, the climbing fern (Lygodium palmatum) of eastern North America will only grow in moist, intensely acid soils, while the bulblet bladder fern (Cystopteris bulbifera), with an overlapping range, is only 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.[24]

Life cycle

Onoclea sensibilis 3 crop

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 they lack 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:

  1. A diploid sporophyte phase produces haploid spores by meiosis (a process of cell division which reduces the number of chromosomes by a half).
  2. 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.
  5. The fertilized egg is now a diploid zygote and grows by mitosis into a diploid sporophyte (the typical fern plant).


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 by some tropical persons (for example in budu pakis, a traditional dish of Brunei[25]) 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.[26][27] Fern tubers were used by the Guanches to make gofio in the Canary Islands. Ferns are generally not known to be poisonous to humans.[28] Licorice fern rhizomes were chewed by the natives of the Pacific Northwest for their flavor.

Ferns of the genus Azolla are very small, floating plants that do not resemble ferns. Called the mosquito fern, they 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.[1]

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 are noxious weeds or invasive species, including Japanese climbing fern (Lygodium japonicum), mosquito fern and sensitive fern (Onoclea sensibilis). Giant water fern (Salvinia molesta) is one of the world's worst aquatic weeds. The important fossil fuel coal consists of the remains of primitive plants, including ferns.

Ferns have been studied and found to be useful in the removal of heavy metals, especially arsenic, from the soil. Other ferns with some economic significance include:


Nature print, Alois Auer


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 is a term for the Victorian era craze of 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.[29]

Barnsley fern plotted with VisSim


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


Ferns figure in folklore, for example in legends about mythical flowers or seeds.[30] 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.[31]

Organisms confused with ferns


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

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.



Fern leaf, probably Blechnum nudum

Tree Fern

A tree fern unrolling a new frond


Tree fern, probably Dicksonia antarctica


Tree ferns, probably Dicksonia antarctica

Haeckel Filicinae 92

"Filicinae" from Ernst Haeckel's Kunstformen der Natur, 1904

Oaxaca fern

Unidentified tree fern in Oaxaca

Tree Fern Spores

Tree Fern Spores San Diego, CA


Leaf of fern

Fern leaves

Unidentified fern with spores showing in Rotorua, NZ.


Ferns in one of many natural Coast Redwood undergrowth settings Santa Cruz, CA.

Asplenium marinum Moore38

Nature prints in The Ferns of Great Britain and Ireland used fronds to produce the plates

Spiraled fern

A young, newly formed fern frond


Fern bed under a forest canopy in woods near Franklin, Virginia

Pyrrosia piloselloides duit-duit

Pyrrosia piloselloides, Dragon's Scale, in Malaysia

Samambaia fern

Fern growing on a wall

Fern Spores

Spores of Dryopteris filix-mas

See also


  1. ^ President, International Association of Pteridologists


  1. ^ Wattieza, Stein, W. E.; Mannolini, F.; Hernick, L. V.; Landling, E.; Berry, C. M. (2007). "Giant cladoxylopsid trees resolve the enigma of the Earth's earliest forest stumps at Gilboa". Nature. 446 (7138): 904–907. Bibcode:2007Natur.446..904S. doi:10.1038/nature05705. PMID 17443185.
  2. ^ a b c d Pteridophyte Phylogeny Group 2016.
  3. ^ McCausland, Jim (20 September 2009). "Rediscover ferns". Sunset.com. Retrieved 2013-09-07.
  5. ^ "Pteridopsida: Fossil Record". University of California Museum of Paleontology. Retrieved 2014-03-11.
  6. ^ Bomfleur B, McLoughlin S, Vajda V (March 2014). "Fossilized nuclei and chromosomes reveal 180 million years of genomic stasis in royal ferns". Science. 343 (6177): 1376–7. Bibcode:2014Sci...343.1376B. doi:10.1126/science.1249884. PMID 24653037.
  7. ^ Large, Mark F.; Braggins, John E. (2004). Tree Ferns. Timber Press. ISBN 0881926302.
  8. ^ "Fern Fronds". Basic Biology. Retrieved 2014-12-06.
  9. ^ a b c Christenhusz & Chase 2014.
  10. ^ a b Smith et al. 2006.
  11. ^ Stace, Clive (2010b). New Flora of the British Isles (3rd ed.). Cambridge, UK: Cambridge University Press. p. xxviii. ISBN 978-0-521-70772-5.
  12. ^ Rai, Hardeep S. & Graham, Sean W. (2010). "Utility of a large, multigene plastid data set in inferring higher-order relationships in ferns and relatives (monilophytes)". American Journal of Botany. 97 (9): 1444–1456. doi:10.3732/ajb.0900305. PMID 21616899., p. 1450
  13. ^ a b c Grewe, Felix; et al. (2013). "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". BMC Evolutionary Biology. 13 (1): 1–16. doi:10.1186/1471-2148-13-8. ISSN 1471-2148. PMC 3553075Freely accessible. PMID 23311954. Retrieved 2013-05-21.
  14. ^ Cantino et al 2007.
  15. ^ a b Chase & Reveal 2009.
  16. ^ a b Smith et al.2006.
  17. ^ Schuettpelz 2007.
  18. ^ Karol, Kenneth G; et al. (2010). "Complete plastome sequences of Equisetum arvense and Isoetes flaccida: implications for phylogeny and plastid genome evolution of early land plant lineages". BMC Evolutionary Biology. 10 (1): 321–336. doi:10.1186/1471-2148-10-321. ISSN 1471-2148. PMC 3087542Freely accessible. PMID 20969798. Retrieved 2013-05-21.
  19. ^ Li, F-W; Kuo, L-Y; Rothfels, CJ; Ebihara, A; Chiou, W-L; et al. (2011). "rbcL and matK Earn Two Thumbs Up as the Core DNA Barcode for Ferns". PLoS ONE. 6 (10): e26597. Bibcode:2011PLoSO...626597L. doi:10.1371/journal.pone.0026597. PMC 3197659Freely accessible. PMID 22028918.
  20. ^ Schneider et al 2009.
  21. ^ Christenhusz et al 2011.
  22. ^ Schuettpelz, Eric. "Fern Phylogeny Inferred from 400 Leptosporangiate Species and Three Plastid Genes," contained in "The Evolution and Diversification of Epiphytic Ferns." Doctoral dissertation, Duke University. 2007. http://dukespace.lib.duke.edu/dspace/bitstream/10161/181/1/D_Schuettpelz_Eric_a_052007.pdf
  23. ^ Ferns Brimming With Life
  24. ^ Walker, Matt (19 February 2010). "A mouse that eats ferns like a dinosaur". BBC Earth News. Retrieved 2010-02-20.
  25. ^ Indigenous Fermented Foods of Southeast Asia. 2015.
  26. ^ "Stone Age humans liked their burgers in a bun", Sonia Van Gilder Cooke, New Scientist, 23 October 2010, p. 18.
  27. ^ "Thirty thousand-year-old evidence of plant food processing" by Anna Revedin et al., PNAS, published online 18 October 2010.
  28. ^ Pelton, Robert (2011). The Official Pocket Edible Plant Survival Manual. Freedom and Liberty Foundation Press. p. 25. BNID 2940013382145.
  29. ^ Boyd, Peter D. A. (2 January 2002). "Pteridomania - the Victorian passion for ferns". Revised: web version. Antique Collecting 28, 6, 9–12. Retrieved 2007-10-02.
  30. ^ May, Lenore Wile (1978). "The economic uses and associated folklore of ferns and fern allies". The Botanical Review. 44 (4): 491–528. doi:10.1007/BF02860848.
  31. ^ "Traditional Finnish Midsummer celebration". Saunalahti.fi. Retrieved 2013-09-07.


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