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Ferns first appear in the fossil record about 360 million years ago in the Kingdom: Plantae
middle Devonian period, but many of the current families and species did Clade: Tracheophytes
not appear until roughly 145 million years ago in the early Cretaceous,
after flowering plants came to dominate many environments. The fern Division: Polypodiophyta
Osmunda claytoniana is a paramount example of evolutionary stasis; Class: Polypodiopsida
paleontological evidence indicates it has remained unchanged, even at the
Cronquist, Takht. &
level of fossilized nuclei and chromosomes, for at least 180 million years.
W.Zimm.
Ferns are not of major economic importance, but some are used for food,
medicine, as biofertilizer, as ornamental plants and for remediating Subclasses[2]
contaminated soil. They have been the subject of research for their ability
to remove some chemical pollutants from the atmosphere. Some fern Extant
species, such as bracken (Pteridium aquilinum) and water fern (Azolla
filiculoides) are significant weeds worldwide. Some fern genera, such as Equisetidae
Azolla, can fix nitrogen and make a significant input to the nitrogen Marattiidae
nutrition of rice paddies. They also play certain roles in folklore.
Ophioglossidae
Polypodiidae
Contents Extinct
Description †Cladoxylopsida
Taxonomy †Zygopteridales
Molecular phylogenetics
†Stauropteridales
Phylogeny
Nomenclature and subdivision †Rhacophytales
Evolution and biogeography Synonyms
Distribution and habitat
Ecology Monilophyta
Life cycle Polypodiophyta
Uses Filicophyta
Culture Filices
Pteridologist
Pteridomania
Folklore
Organisms confused with ferns
Misnomers
Fern-like flowering plants
Gallery
See also
Notes
References
Bibliography
Books and theses
Journal articles
Websites
External links
Description
Like the sporophytes of seed plants, those of ferns consist of stems, leaves
and roots. Ferns differ from seed plants in reproducing by spores. 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.
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 Ferns at the Royal Melbourne
terrestrial ones have above-ground creeping stolons (e.g., Polypodiaceae), Botanical Gardens
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).[3]
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:
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:
Antheridia: Small spherical structures that produce flagellate sperm.
Archegonia: A flask-shaped structure that produces a single egg at the bottom, reached by the
sperm by swimming down the neck.
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.
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).[6][7][8] By 1806 this had increased to 38 genera,[9] and
has progressively increased since (see Schuettpelz et al (2018) Figure 1). 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.[10] 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:[11]
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.[11] However, there remained differences of opinion about
the placement of the genus Equisetum (see Equisetopsida for further discussion). One possible solution was to
denominate only 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".[12] 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".[13] Grewe
et al. (2013) confirmed the inclusion of horsetails within ferns sensu lato, but also suggested that uncertainties
remained in their precise placement.[14] Other classifications have raised Ophioglossales to the rank of a fifth class,
separating the whisk ferns and ophioglossoid ferns.[14]
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.
Phylogeny
The ferns are related to other higher order taxa, as shown in the following cladogram:[10][15][16][2]
Polypodiophyta
(Polypodiopsida) - ferns
Tracheophyta - vascular plants
Euphyllophyta Gymnospermae
Spermatophyta - seed plants
Angiospermae - flowering
plants
In addition they defined 11 orders and 37 families.[11] That system was a consensus of a number of studies, and was
further refined.[14][18] The phylogenetic relationships are shown in the following cladogram (to the level of
orders).[11][19][14] This division into four major clades was then confirmed using morphology alone.[20]
Tracheophyta
Lycopodiophytes (club mosses, spike mosses, quillworts)
Euphyllophytes Spermatophytes (seed plants)
Equisetopsida
Equisetales (horsetails)
Marattiopsida Marattiales
Leptosporangiate
Polypodiopsida Ferns
Osmundales
Gleicheniales
Schizaeales
Salviniales (heterosporous)
Polypodiales
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).[16] 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;[10]
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]
Equisetidae
Ophioglossidae
Polypodiopsida
Marattiidae
Polypodiidae
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.[22] 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.)[2] 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.[8][23]
Smith et al. Chase & Reveal Christenhusz et al. Christenhusz & Chase
PPG I (2016)[2]
(2006)[11] (2009)[16] (2011)[21] (2014, 2018)[10][24]
ferns
ferns monilophytes ferns (Polypodiophyta) Class
(monilophytes)
(no rank) (no rank) (no rank) Polypodiopsida
(no rank)
Class Subclass Subclass Subclass
Subclass Equisetidae
Equisetopsida Equisetidae Equisetidae Equisetidae
Subclass
Class Ophioglossidae Subclass Subclass
Subclass Ophioglossidae
Psilotopsida Subclass Ophioglossidae Ophioglossidae
Psilotidae
Class Subclass Subclass Subclass
Subclass Marattiidae
Marattiopsida Marattiidae Marattiidae Marattiidae
Class Subclass Subclass Subclass
Subclass Polypodiidae
Polypodiopsida Polypodiidae Polypodiidae Polypodiidae
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.[25][1][26][27] Ferns evolved to cope with low-light
conditions present under the canopy of angiosperms. Remarkably, the fern photoreceptor neochrome was obtained
via horizontal gene transfer from a bryophyte lineage.[28]
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.[32]
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.[33]
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
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
Gametophyte (thalloid green mass)
sporophyte.
and sporophyte (ascendent frond) of
The life cycle of a typical fern proceeds as follows: 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[34]) 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.[35][36] Fern tubers were used by the Guanches to make gofio in the Canary Islands. Ferns are generally not
known to be poisonous to humans.[37] Licorice fern rhizomes were chewed by the natives of the Pacific Northwest
for their flavor.
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.[38]
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[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] 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:
Dryopteris filix-mas (male fern), used as a vermifuge, and formerly in the US Pharmacopeia; also, this
fern accidentally sprouting in a bottle resulted in Nathaniel Bagshaw Ward's 1829 invention of the
terrarium or Wardian case
Rumohra adiantiformis (floral fern), extensively used in the florist trade
Microsorum pteropus (Java fern), one of the most popular freshwater aquarium plants.
Osmunda regalis (royal fern) and Osmunda cinnamomea (cinnamon fern), the root fiber being used
horticulturally; the fiddleheads of O. cinnamomea are also used as a cooked vegetable
Matteuccia struthiopteris (ostrich fern), the fiddleheads used as a cooked vegetable in North America
Pteridium aquilinum and Pteridium esculentum (bracken), the fiddleheads used as a cooked
vegetable in Japan and are believed to be responsible for the high rate of stomach cancer in Japan. It
is also one of the world's most important agricultural weeds, especially in the British highlands, and
often poisons cattle and horses.
Diplazium esculentum (vegetable fern), a source of food for some societies
Pteris vittata (brake fern), used to absorb arsenic from the soil
Polypodium glycyrrhiza (licorice fern), roots chewed for their pleasant flavor
Tree ferns, used as building material in some tropical areas
Cyathea cooperi (Australian tree fern), an important invasive species in Hawaii
Ceratopteris richardii, a model plant for teaching and research, often called C-fern
Culture
Pteridologist
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 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.[42]
The dried form of ferns was also used in other arts, being used 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.
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 hydrozoan 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.
Gallery
A tree fern unrolling Tree fern, probably Tree ferns, probably "Filicinae" from
a new frond Dicksonia antarctica Dicksonia antarctica Ernst Haeckel's
Kunstformen der
Natur, 1904
See also
British Pteridological Society
Chirosia betuleti - Fern gall
Fern spike
Fern sports
Paisley (design)
Pteridophyte
Silver fern flag
Notes
a. President, International Association of Pteridologists
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External links
Data related to Pteridophyta at Wikispecies
Media related to Polypodiopsida at Wikimedia Commons
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