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''Asplenium'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of about 700
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of
fern The ferns (Polypodiopsida or Polypodiophyta) are 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 tissue ...
s, often treated as the only genus in the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Aspleniaceae, though other authors consider '' Hymenasplenium'' separate, based on
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
analysis of
DNA sequence A nucleic acid sequence is a succession of bases within the nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. This succession is denoted by a series of a set of five different letters that indicate the order of the nu ...
s, a different chromosome count, and structural differences in the rhizomes. The
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
for the genus is '' Asplenium marinum''. The most common vernacular name is spleenworts, applied to the more "typical" species. '' A. nidus'' and several similar species are called bird's-nest ferns, the ''Camptosorus'' group is known as walking ferns, and distinct names are applied to some other particularly well-known species.


Etymology

From
New Latin Neo-LatinSidwell, Keith ''Classical Latin-Medieval Latin-Neo Latin'' in ; others, throughout. (also known as New Latin and Modern Latin) is the style of written Latin used in original literary, scholarly, and scientific works, first in Italy d ...
''asplenium'' (“spleenworts”),
Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
's adjustment of
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
''asplenon'' (“spleenwort”), from
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
ἄσπληνον (''ásplēnon'', “spleenwort”), from ἀ- (''a-'', “un-”) + σπλήν (''splḗn'', “spleen”) + -ον (''-on'', “-um”), from its use to cure anthrax in livestock.


Taxonomy and genetics

Many groups of species have been separated from ''Asplenium'' as segregate genera. These include ''Camptosorus'', ''Ceterach'', ''Phyllitis'', and ''Tarachia'', but these species can form hybrids with other ''Asplenium'' species and because of this are usually included in a more broadly defined ''Asplenium''. Some of the older classifications elevate the Aspleniaceae to the
taxonomic rank In biology, taxonomic rank (which some authors prefer to call nomenclatural rank because ranking is part of nomenclature rather than taxonomy proper, according to some definitions of these terms) is the relative or absolute level of a group of or ...
of
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
as Aspleniales. The newer classifications place it in the subordinal group called eupolypods within the order
Polypodiales The Order (biology), order Polypodiales encompasses the major lineages of polypod ferns, which comprise more than 80% of today's fern species. They are found in many parts of the world including Tropics, tropical, semitropical and Temperate clima ...
. Within the eupolypods, Aspleniaceae belongs to a
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
informally and provisionally known as eupolypods II. It has been found that in some species, the
chloroplast A chloroplast () is a type of membrane-bound organelle, organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. Chloroplasts have a high concentration of chlorophyll pigments which captur ...
genome A genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as ...
has
evolved Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
in complex and highly unusual ways. This makes standard
cladistic Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...
analyses unsuited to resolve the
phylogeny A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...
of that particular group of ferns, and even very sophisticated
computational phylogenetics Computational phylogenetics, phylogeny inference, or phylogenetic inference focuses on computational and optimization algorithms, Heuristic (computer science), heuristics, and approaches involved in Phylogenetics, phylogenetic analyses. The goal i ...
methods yield little information. In addition to hybridization running rampant in parts of this genus, there are also some species like the mother spleenwort (''A. bulbiferum'') or '' A. viviparum'' which mainly reproduce asexually, essentially
cloning Cloning is the process of producing individual organisms with identical genomes, either by natural or artificial means. In nature, some organisms produce clones through asexual reproduction; this reproduction of an organism by itself without ...
themselves over and over again. While most are
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
or
tetraploid Polyploidy is a condition in which the cells of an organism have more than two paired sets of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two complete sets of chromosomes, one fro ...
, some species (e.g. '' A. shuttleworthianum'') are octoploid.


Uses

Both the scientific name and the common name "spleenwort" are derived from an old belief, based on the doctrine of signatures, that the fern was useful for ailments of the
spleen The spleen (, from Ancient Greek '' σπλήν'', splḗn) is an organ (biology), organ found in almost all vertebrates. Similar in structure to a large lymph node, it acts primarily as a blood filter. The spleen plays important roles in reg ...
, due to the spleen-shaped sori on the backs of the
frond A frond is a large, divided leaf. In both common usage and botanical nomenclature, the leaves of ferns are referred to as fronds and some botanists restrict the term to this group. Other botanists allow the term frond to also apply to the lar ...
s. "-wort" is an ancient English term that simply means "
plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...
" (compare German ''-wurz''). The plants were thought to cause
infertility In biology, infertility is the inability of a male and female organism to Sexual reproduction, reproduce. It is usually not the natural state of a healthy organism that has reached sexual maturity, so children who have not undergone puberty, whi ...
in women.
Vitruvius Vitruvius ( ; ; –70 BC – after ) was a Roman architect and engineer during the 1st century BC, known for his multi-volume work titled . As the only treatise on architecture to survive from antiquity, it has been regarded since the Renaissan ...
relates the story of the name thus:
... certain pastures in Crete, on each side of the river Pothereus, which separates the two Cretan states of Gnosus and Gortyna. There are cattle at pasture on the right and left banks of that river, but while the cattle that feed near Gnosus have the usual spleen, those on the other side near Gortyna have no perceptible spleen. On investigating the subject, physicians discovered on this side a kind of herb which the cattle chew and thus make their spleen small. The herb is therefore gathered and used as a medicine for the cure of splenetic people. The Cretans call it ἄσπληνον.
A few of these ferns have some economic importance in the
horticulture Horticulture (from ) is the art and science of growing fruits, vegetables, flowers, trees, shrubs and ornamental plants. Horticulture is commonly associated with the more professional and technical aspects of plant cultivation on a smaller and mo ...
trade. The bird's-nest ferns ('' A. nidus'' and several very similar, closely related species) are commonly found for sale as a house plant. The Australian mother spleenwort (''A. bulbiferum'') is sometimes available at
greenhouse A greenhouse is a structure that is designed to regulate the temperature and humidity of the environment inside. There are different types of greenhouses, but they all have large areas covered with transparent materials that let sunlight pass an ...
s, and is of interest, along with the related '' A. viviparum'', for the many small bulblets borne on the fronds that may grow into new plants. This characteristic is also shared with the eastern North American walking fern (''A. rhizophyllum'') and several Mexican species including '' A. palmeri''. The ebony spleenwort ''A. platyneuron'' is also sometimes sold in nurseries as a hardy plant. However, many spleenworts are epipetric or
epiphytic An epiphyte is a plant or plant-like organism that grows on the surface of another plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphyt ...
and difficult to cultivate. ''Asplenium'' species are used as food plants by the
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e of some
Lepidoptera Lepidoptera ( ) or lepidopterans is an order (biology), order of winged insects which includes butterflies and moths. About 180,000 species of the Lepidoptera have been described, representing 10% of the total described species of living organ ...
species including '' Batrachedra bedelliella'' which feeds exclusively on ''A. nidus''. For diseases of ''Asplenium'', see List of foliage plant diseases (Polypodiaceae).


Selected species

* '' Asplenium adiantum-nigrum'' L.black spleenwort (= ''A. lucidum'' Burm.f.) ** ''Asplenium adiantum-nigrum'' ssp. ''adiantum-nigrum'' ** ''Asplenium adiantum-nigrum'' ssp. ''serpentini'' (Tausch) Koch * '' Asplenium adulterinum'' Mildeladder spleenwort * '' Asplenium aequibasis'' ( C.Chr.) J.P.Roux * '' Asplenium aethiopicum'' ( Burm.f.) Bech. * '' Asplenium africanum'' * '' Asplenium × alternifolium'' Wulf. * '' Asplenium anceps'' owe ex Hook. et Grev. Page (1971)/small> * '' Asplenium angustum'' Sw. * '' Asplenium antiquum'' Makino * '' Asplenium arcanum'' A.R.Sm. * '' Asplenium ascensionis'' S.Watson * '' Asplenium attenuatum'' Sw. * '' Asplenium aureum'' (sometimes in ''Ceterach'') * '' Asplenium auritum'' R.Br. * '' Asplenium australasicum'' (J.Sm.) Hook.crow's-nest fern ** ''Asplenium australasicum'' f. ''australasicum'' ** ''Asplenium australasicum'' f. ''robinsonii'' * '' Asplenium azoricum'' Lovis, Rasbach & Reichst. * '' Asplenium bifrons'' Sodiro * '' Asplenium billottii'' – lanceolate spleenwort * '' Asplenium bipinnatifidum'' * '' Asplenium brachycarpum'' * '' Asplenium bradleyi'' * '' Asplenium bulbiferum'' – mother spleenwort, hen and chickens fern, ''mouku'' ( Māori) * '' Asplenium carnarvonense'' - Brownsey * '' Asplenium caudatum'' * '' Asplenium ceterach'' – rustyback fern (sometimes in ''Ceterach'') * '' Asplenium chathamense'' Brownsey * '' Asplenium chihuahuense'' Baker * '' Asplenium compressum'' Sw. * '' Asplenium congestum'' C.Chr. * ''
Asplenium corderoanum ''Asplenium'' is a genus of about 700 species of fern The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from ...
'' * '' Asplenium crinicaule'' * '' Asplenium cristatum'' * '' Asplenium cuneifolium'' Viv. (= ''A. forsteri'' ''auct. non'' Sadl.) – serpentine spleenwort * '' Asplenium cymbifolium'' * '' Asplenium daghestanicum'' H.Christ – Dagestanian spleenwort * '' Asplenium dalhousiae'' (sometimes in ''Ceterach'') * '' Asplenium dareoides'' * '' Asplenium daucifolium'' Lam.Mauritius spleenwort * '' Asplenium difforme'' R.Br. * '' Asplenium fissum'' * '' Asplenium dimorphum''
Kunze Kunz, Künz, or Kunze is a surname. Notable people with the surname include: *Kunz (singer) (Marco Kunz, born 1985), Swiss singer *Adrian Kunz (born 1967), Swiss international footballer *Alfred Kunz (Catholic priest) (1931–1998), American murder ...
Norfolk Island Spleenwort * '' Asplenium divaricatum'' * ''
Asplenium dregeanum ''Asplenium'' is a genus of about 700 species of ferns, often treated as the only genus in the family Aspleniaceae, though other authors consider '' Hymenasplenium'' separate, based on molecular phylogenetic analysis of DNA sequences, a differen ...
'' * '' Asplenium × ebenoides'' R.R.Scott * '' Asplenium ecuadorense'' Stolze * '' Asplenium feei'' Kunze ex Fée * '' Asplenium fissum'' * '' Asplenium flabellifolium'' Cav.necklace fern * '' Asplenium flaccidum'' G.Forst.weeping spleenwort, hanging spleenwort * '' Asplenium fontanum'' (L.) Bernh.smooth rock spleenwort * '' Asplenium forisiense'' – rock spleenwort * '' Asplenium formosum'' * '' Asplenium gemmiferum'' Schrad. * '' Asplenium × germanicum'' * '' Asplenium gueinzii'' Mett. * '' Asplenium goudeyi'' Lord Howe Island *'' Asplenium haughtonii'' – Barn Fern * '' Asplenium hemionitis'' * '' Asplenium hermannii-christii'' Fomin – Hermann Christ's asplenium * '' Asplenium hookerianum'' Colenso * '' Asplenium hybridum'' * '' Asplenium incisum'' * '' Asplenium × jacksonii'' Alston – Jackson's spleenwort (sterile, triploid hybrid between '' Asplenium adiantum-nigrum'' and '' Asplenium scolopendrium'') * '' Asplenium × kenzoi'' - oni-hinokishida, cultivated in Japan * '' Asplenium komarovii'' - Akasawa * '' Asplenium laciniatum'' * '' Asplenium lamprophyllum'' Carse * '' Asplenium laserpitiifolium'' – Johnston River fern * '' Asplenium lepidum'' C.Presl * '' Asplenium listeri'' – Christmas Island spleenwort * '' Asplenium longissimum'' * '' Asplenium lucidum'' * '' Asplenium lunulatum'' – Hen-and-chicks * '' Asplenium lyallii'' * '' Asplenium macedonicum'' * '' Asplenium majoricum'' * '' Asplenium marinum'' – sea spleenwort * '' Asplenium × microdon'' T Moore – Moore's spleenwort (hybrid between ''Asplenium scolopendrium'' and ''Asplenium obovatum'' subsp ''lanceolatum'') * '' Asplenium milnei'' Carruth * '' Asplenium montanum'' – mountain spleenwort * '' Asplenium musifolium'' * '' Asplenium nidus'' – bird's-nest fern * '' Asplenium normale'' * '' Asplenium obliquum'' * '' Asplenium oblongifolium'' Colensoshining spleenwort (= ''A. lucidum'' ''auct. non'' Burm.f., ''sensu'' G.Forst.) * '' Asplenium obovatum'' * '' Asplenium obtusatum'' G.Forst. ** ''Asplenium obtusatum'' ssp. ''northlandicum'' (Brownsey) Ogle (possibly distinct species) ** ''Asplenium obtusatum'' 'Chile' (possibly distinct species, sometimes included in ''A. obliquum'') * '' Asplenium oligolepidum'' C.Chr. (= ''A. lucidum'' ''auct. non'' Burm.f., ''sensu'' G.Forst.) * '' Asplenium oligophlebium'' * '' Asplenium onopteris'' L.western black spleenwort, Irish spleenwort (sometimes included in ''A. adiantum-nigrum'') * '' Asplenium pacificum'' * '' Asplenium paleaceum'' R.Br.chaffy spleenwort * '' Asplenium palmeri'' * '' Asplenium parvum'' * '' Asplenium petrarchae'' * '' Asplenium pinnatifidum'' – lobed spleenwort * '' Asplenium planicaule'' * '' Asplenium platybasis'' Kunze ex Mett. * '' Asplenium platyneuron'' – ebony spleenwort * '' Asplenium polyodon'' G.Forst.sickle spleenwort * '' Asplenium praemorsum'' * '' Asplenium prolongatum'' Hook. * '' Asplenium pteridoides'' Baker * '' Asplenium resiliens'' – black-stemmed spleenwort * '' Asplenium rhizophyllum'' – American walking fern (sometimes in ''Camptosorus'') * '' Asplenium richardii'' * '' Asplenium ruprechtii'' – Asian walking fern (sometimes in ''Camptosorus'') * '' Asplenium ruta-muraria'' L.wall-rue * '' Asplenium rutifolium'' * '' Asplenium sagittatum'' – Mule's spleenwort (sometimes in ''Phyllitis'') * '' Asplenium sandersonii'' Hook. * '' Asplenium × sarniense'' Sleep Guernsey Spleenwort * '' Asplenium schizotrichum'' Copel. * '' Asplenium schweinfurthii'' * '' Asplenium scleroprium'' * '' Asplenium scolopendrium'' – hart's-tongue fern (sometimes in ''Phyllitis'') ** ''Asplenium scolopendrium'' var. ''americanum'' – American Hart's-tongue * '' Asplenium seelosii'' * '' Asplenium septentrionale'' – forked spleenwort, northern spleenwort * '' Asplenium septentrionale × trichomanes'' Wulf. * '' Asplenium serra'' * '' Asplenium serratum'' – wild bird's-nest fern * '' Asplenium sessilifolium'' * '' Asplenium shuttleworthianum'' Kunze * '' Asplenium simplicifrons'' F.Muell. * '' Asplenium splendens'' * '' Asplenium surrogatum'' P.S.Green * '' Asplenium tenerum'' G.Forst. * '' Asplenium terrestre'' * '' Asplenium theciferum'' (Kunth) Mett. * '' Asplenium thunbergii'' * ''
Asplenium trichomanes ''Asplenium trichomanes'', the maidenhair spleenwort (not to be confused with the similar-looking Adiantum, maidenhair fern), is a small fern in the spleenwort genus ''Asplenium''. It is a widespread and common species, occurring almost worldwide ...
'' – maidenhair spleenwort ** ''Asplenium trichomanes'' ssp. ''quadrivalens'' D.E. Meyer ** ''Asplenium trichomanes'' ssp. ''trichomanes'' ** ''Asplenium trichomanes'' subsp. ''coriaceifolium'' * '' Asplenium trilobum ''Cav. * '' Asplenium tutwilerae'' B.R.Keener & L.J.Davenport * '' Asplenium unisorum'' (Wagner) Viane * '' Asplenium vespertinum'' * '' Asplenium vieillardii'' Mett. * '' Asplenium virens'' * '' Asplenium viride'' – green spleenwort * '' Asplenium vittiforme'' * '' Asplenium viviparum'' * '' Asplenium yoshinagae'' Makino


See also

* Mount Asplenium * '' Phyllocladus aspleniifolius'' (Celery-top Pine, a
conifer Conifers () are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a sin ...
with ''Asplenium''-like leaves) * ''Asplenium'' hybrids


References

{{Authority control Epiphytes Fern genera Taxa named by Carl Linnaeus