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The Aspleniaceae (spleenworts) are a family of
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except th ...
s, included in the order
Polypodiales The 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 tropical, semitropical and temperate areas. Description Polypodiale ...
. The composition and classification of the family have been subject to considerable changes. In particular, there is a narrow
circumscription Circumscription may refer to: *Circumscribed circle *Circumscription (logic) *Circumscription (taxonomy) * Circumscription theory, a theory about the origins of the political state in the history of human evolution proposed by the American anthrop ...
, Aspleniaceae s.s. (adopted here), in which the family contains only two genera, and a very broad one, Aspleniaceae
s.l. SL may refer to: Arts and entertainment * SL (rapper), a rapper from London * ''Second Life'', a multi-user 3D virtual world * Sensei's Library, an Internet site dedicated to the game of Go * Subdominant leittonwechselklänge * Leica SL, a mirr ...
, in which the family includes 10 other families kept separate in the narrow circumscription, with the Aspleniaceae s.s. being reduced to the subfamily Asplenioideae. The family has a worldwide distribution, with many species in both temperate and tropical areas. Elongated unpaired sori are an important characteristic of most members of the family.


Description

Members of the family grow from
rhizome In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow ho ...
s, that are either creeping or somewhat erect, and are usually but not always unbranched, and have scales that usually have a lattice-like (clathrate) structure. In some species, for example ''
Asplenium nidus :''See bird's-nest fern for other plants with this common name.'' ''Asplenium nidus'' is an epiphytic species of fern in the family Aspleniaceae, native to tropical southeastern Asia, eastern Australia, Hawaii (''ʻēkaha'' in Hawaiian), Po ...
'', the rhizomes form a kind of basket which collects detritus. The leaves may be undivided or be divided, with up to four-fold
pinnation Pinnation (also called pennation) is the arrangement of feather-like or multi-divided features arising from both sides of a common axis. Pinnation occurs in biological morphology, in crystals, such as some forms of ice or metal crystals, and in ...
. The sori are characteristic of the family. They are elongated, and normally located on one side of a vein. More rarely, they may be in pairs on a single vein, but then they never curve over the vein. A flap-like
indusium A sorus (pl. sori) is a cluster of sporangia (structures producing and containing spores) in ferns and fungi. A coenosorus (plural coenosori) is a compound sorus composed of multiple, fused sori. Etymology This New Latin word is from Ancient G ...
arises along one edge of a sorus. The leaf stalks ( petioles) have two
vascular bundle A vascular bundle is a part of the transport system in vascular plants. The transport itself happens in the stem, which exists in two forms: xylem and phloem. Both these tissues are present in a vascular bundle, which in addition will includ ...
s, uniting to form an X-shape in cross-section towards the tip of the leaf. The stalks of the
sporangia A sporangium (; from Late Latin, ) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their lif ...
are one cell wide in the middle.


Taxonomy

The family Aspleniaceae was first described by Edward Newman in 1840. Newman included three genera: ''
Athyrium ''Athyrium'' (lady-fern) is a genus of about 180 species of terrestrial ferns, with a cosmopolitan distribution. It is placed in the family Athyriaceae, in the order Polypodiales. Its genus name is from Greek '' a-'' ('without') and Latinized G ...
'', ''
Asplenium ''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 different ...
'' and ''Scolopendrium''. ''Athyrium'' is now placed in a different family,
Athyriaceae The Athyriaceae (ladyferns and allies) are a family of terrestrial ferns in the order Polypodiales. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family is placed in the suborder Aspleniineae, and includes two ge ...
, not considered very strongly related to the Aspleniaceae, and ''Scolopendrium'' is regarded as synonym of ''Asplenium''. The narrow
circumscription Circumscription may refer to: *Circumscribed circle *Circumscription (logic) *Circumscription (taxonomy) * Circumscription theory, a theory about the origins of the political state in the history of human evolution proposed by the American anthrop ...
of the family adopted by the
Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group, or PPG, is an informal international group of systematic botanists who collaborate to establish a consensus on the classification of pteridophytes (lycophytes and ferns) that reflects knowledge about plant relation ...
classification of 2016 (PPG I) recognizes only two genera, ''
Asplenium ''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 different ...
'' and ''
Hymenasplenium ''Hymenasplenium'' is one of three genera of ferns in the Aspleniaceae (spleenwort family), in the eupolypods II clade of the order Polypodiales. The others are '' Hemidictyum'' and ''Asplenium''. ''Hymenasplenium'' was segregated because it is ...
''. ''Asplenium'' has previously been split into a dozen or so genera, including ''Diella'', found only in Hawaii. The consensus of
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 ...
studies is that all are nested within ''Asplenium''. PPG I places Aspleniaceae in the suborder
Aspleniineae Aspleniineae is a suborder of ferns in the order Polypodiales. It is equivalent to the clade eupolypods II in earlier systems; it is also treated as a single very broadly defined family Aspleniaceae. The suborder generally corresponds with the o ...
of the order
Polypodiales The 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 tropical, semitropical and temperate areas. Description Polypodiale ...
. Earlier, Christenhusz and Chase had proposed a much broader circumscription of Aspleniaceae, in which it consisted of all the separate families that PPG I places in the suborder Aspleniineae (eight at the time), with the families reduced to subfamilies. Thus the Aspleniaceae of PPG I became the subfamily Asplenioideae. , the broader circumscription of the Aspleniaceae is used by
Plants of the World Online Plants of the World Online (POWO) is an online database published by the Royal Botanic Gardens, Kew. It was launched in March 2017 with the ultimate aim being "to enable users to access information on all the world's known seed-bearing plants by ...
, which lists 24 genera.


Phylogenic relationships

Aspleniaceae is placed in a clade known as eupolypods II, or more formally as suborder
Aspleniineae Aspleniineae is a suborder of ferns in the order Polypodiales. It is equivalent to the clade eupolypods II in earlier systems; it is also treated as a single very broadly defined family Aspleniaceae. The suborder generally corresponds with the o ...
. The following cladogram, based on Lehtonen (2011) and Rothfels & al. (2012), shows a likely
phylogenic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
relationship between the Aspleniaceae and the other families in the clade.


Genera

In the PPG I system, Aspleniaceae s.s. contains two genera: *''
Asplenium ''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 different ...
'' L. – about 700 species, worldwide *''
Hymenasplenium ''Hymenasplenium'' is one of three genera of ferns in the Aspleniaceae (spleenwort family), in the eupolypods II clade of the order Polypodiales. The others are '' Hemidictyum'' and ''Asplenium''. ''Hymenasplenium'' was segregated because it is ...
'' Hayata – about 40 species, tropical and subtropical


Distribution and habitat

The Aspleniaceae have a worldwide distribution, with the large genus ''Asplenium'' being native to almost all parts of the world except Antarctica and some high Arctic areas. The family is unusual in having high diversity in both temperate and tropical areas, and more-or-less equal numbers of terrestrial and epiphytic species. Plants are terrestrial, growing in the ground,
lithophytic Lithophytes are plants that grow in or on rocks. They can be classified as either epilithic (or epipetric) or endolithic; epilithic lithophytes grow on the surfaces of rocks, while endolithic lithophytes grow in the crevices of rocks (and are also ...
, growing on rocks, or
epiphytic An epiphyte is an organism that grows on the surface of a 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 epiphytes grow are called phoroph ...
, growing on other plants; less often they are aquatic, growing in moving water.


References


Bibliography


Germplasm Resources Information Network: Aspleniaceae
*

{{Taxonbar, from=Q1137402 Fern families Taxa named by Edward Newman