Vittarioideae is a subfamily of the

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 ...

family

Pteridaceae Pteridaceae is a family of ferns in the order Polypodiales, including some 1150 known species in ca 45 genera (depending on taxonomic opinions), divided over five subfamilies. The family includes four groups of genera that are sometimes recogni ...

, in 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 ...

. The subfamily includes the previous families Adiantaceae (adiantoids or maidenhair ferns) and Vittariaceae (vittarioids or shoestring ferns).

Description

The subfamily includes two distinct groups of ferns: the adiantoids, consisting of the single genus ''

Adiantum ''Adiantum'' (), the maidenhair fern (not to be confused with the similar-looking maidenhair spleenwort fern), is a genus of about 250 species of ferns in the subfamily Vittarioideae of the family Pteridaceae, though some researchers place it ...

'', and the vittarioids, several genera, including ''

Vittaria ''Vittaria'', the shoestring ferns, is a genus of ferns in the Vittarioideae subfamily of the family Pteridaceae. It had previously been placed in the family Vittariaceae,Karl U. Kramer. 1990. "Vittariaceae". pages 272-277. In: Klaus Kubitzki ( ...

'', which typically have highly reduced leaves, usually entire, and an epiphytic habit. The ferns historically considered as ''Adiantum'' include both petrophilic and terrestrial plants. The vittarioid ferns are primarily

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 ...

in tropical regions and all have simple leaves with sori that follow the veins and lack true indusia; the sori are most often marginal with a false indusium formed from the reflexed leaf margin. The family also includes a species, ''

Vittaria appalachiana ''Vittaria appalachiana'', or the Appalachian shoestring fern, is a fern species in the subfamily Vittarioideae of the family Pteridaceae. It is native to moist and shaded outcrops in the Appalachian Mountains. It is notable for existing only in ...

'', that is highly unusual in that the

sporophyte A sporophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the biological life cycle, life cycles of plants and algae. It is a diploid multicellular organism which produces asexual Spo ...

stage of the life cycle is absent. This species consists solely of

photosynthetic Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...

gametophyte A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the se ...

s that reproduce asexually.

Taxonomy

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 demonstrated that the vittarioid ferns were nested within the genus ''

'' as it was originally

circumscribed In geometry, a circumscribed circle for a set of points is a circle passing through each of them. Such a circle is said to ''circumscribe'' the points or a polygon formed from them; such a polygon is said to be ''inscribed'' in the circle. * Circum ...

, making that genus paraphyletic. In the

Pteridophyte Phylogeny Group The Pteridophyte Phylogeny Group (PPG) is an informal international group of systematic botanists who collaborate to establish on the classification of pteridophytes ( lycophytes and ferns) that reflects knowledge about plant relationships discove ...

classification of 2016 (PPG I), the family is treated as the subfamily Vittarioideae] of the family

. The following diagram shows a likely phylogenetic relationship between the Vittarioideae and other subfamilies of the Pteridaceae.Schuettpelz ''et al.'' (2007)
Eric Schuettpelz, Harald Schneider, Layne Huiet, Michael D. Windham, Kathleen M. Pryer: "A molecular phylogeny of the fern family Pteridaceae: Assessing overall relationships and the affinities of previously unsampled genera." ''Molecular Phylogenetics and Evolution'' 44 (2007) 1172–1185

History

The first suprageneric classification based on ''Vittaria'' was made by

Carl Borivoj Presl Carl Borivoj Presl (; 17 February 1794 – 2 October 1852) was a Czech botanist. Biography Presl lived his entire life in Prague, and was a professor of botany at the University of Prague (1833–52).tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide use of the term in English is in the discipline of anthropology. The definition is contested, in part due to conflict ...

Vittariaceae to contain the genera ''Vittaria'' and '' Prosaptia'', the latter now included in the grammitid ferns. He invented the new genus ''

Haplopteris ''Haplopteris'' is a genus of vittarioid ferns, a member of subfamily Vittarioideae and family Pteridaceae. Description Like other vittarioids, the members of ''Haplopteris'' are epiphytes. The rhizome has a distinct upper and lower side, lacki ...

'' to accommodate another group of simple-leaved ferns separated from ''

Pteris ''Pteris'' (brake) is a genus of about 300 species of ferns in the subfamily Pteridoideae of the family Pteridaceae. They are native to tropical and subtropical regions, southward to New Zealand, Australia, and South Africa, north to Japan and N ...

'', but placed it in tribe Adiantaceae instead, due to the location of its sori just behind the leaf margin. In his 1911 treatment of the tribe, Ralph Benedict adopted a circumscription similar to modern treatments, within which he recognized the genera ''

Ananthacorus ''Ananthacorus'' is a genus of ferns in the subfamily Vittarioideae of the family Pteridaceae with a single species ''Ananthacorus angustifolius''. Its native distribution ranges from Mexico through Central America to northern South America. It ...

'', ''

Anetium ''Polytaenium'' is a genus of ferns in the subfamily Vittarioideae of the family Pteridaceae. Species are native to Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin ...

'', ''

Antrophyum ''Antrophyum'' is a genus of ferns in the family Pteridaceae.''Antrophyum''
USDA Germplasm Reso ...

'', ''

Hecistopteris ''Hecistopteris'' is a fern genus in the Vittarioideae subfamily of the Pteridaceae. Species The genus ''Hecistopteris'' contains the following species:Monogramma'', ''

Polytaenium ''Polytaenium'' is a genus of ferns in the subfamily Vittarioideae of the family Pteridaceae. Species are native to Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin ...

'', and ''

''. He described ''

Radiovittaria ''Radiovittaria'' is a genus of ferns in the subfamily Vittarioideae of the family Pteridaceae. Species are native to southeast Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost coun ...

'' as a subgenus of ''Vittaria'', subsumed ''

Scoliosorus ''Scoliosorus'' is a genus of ferns in the subfamily Vittarioideae of the family Pteridaceae with a single species, ''Scoliosorus ensiformis''. The species is native to Mexico and Central America Central America is a subregion of North A ...

'' within ''

'' as doubtfully worthy of subgeneric rank, while ''

Rheopteris ''Rheopteris'' is a genus of ferns in the subfamily Vittarioideae of the family Pteridaceae with a single species, ''Rheopteris cheesmaniae'', in the Pteridophyte Phylogeny Group classification of 2016 (PPG I). Other sources sink the genus into ' ...

'' had not yet been discovered. ''Haplopteris'' he explicitly synonymized with ''Vittaria'' in 1914. Carl Christensen used the name "Vittarioideae" in Verdoorn's ''Manual of Pteridology'' in 1938, but did not include a description, leaving it nomenclaturally invalid.

Ren-Chang Ching Ren-Chang Ching (; 15 February 1898 – 22 July 1986), courtesy name Zinong, was a Chinese botanist who specialised in ferns. Life and work Ren-Chang Ching was a Chinese botanist and pteridologist who made significant collections of plants from ...

raised Vittariaceae to the rank of a family in 1940. The first well-sampled

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 ...

study Study or studies may refer to: General * Education **Higher education * Clinical trial * Experiment * Field of study * Observational study * Scientific study * Research * Study skills, abilities and approaches applied to learning Other * Study ...

of the vittarioids was based on 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 ...

gene In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...

rbcL Ribulose-1,5-bisphosphate carboxylase/oxygenase, commonly known by the abbreviations RuBisCo, rubisco, RuBPCase, or RuBPco, is an enzyme () involved in the Photosynthesis#Light-independent reactions, light-independent (or "dark") part of photosyn ...

. In this study, it was found that 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 ...

of ''Monogramma'' is embedded in ''Haplopteris''; the segregation of '' Vaginularia'' from ''Monogramma'' was also supported, as members of ''Vaginularia'' formed a clade sister to ''Rheopteris'' and distant from ''Monogramma'' ''sensu stricto''.Bradley Ruhfel, Stuart Lindsay, and Charles C. Davis. 2008. "Phylogenetic Placement of ''Rheopteris'' and the Polyphyly of ''Monogramma'' (Pteridaceae s.l.): Evidence from rbcL Sequence Data". ''Systematic Botany'' 33(1):37-43, A later molecular phylogeny, published in 2016, established the genus ''

Antrophyopsis ''Antrophyopsis'' is a genus of vittarioid ferns, a member of subfamily Vittarioideae and family Pteridaceae. Like other vittarioids, ferns in the genus are epiphytes with simple, straplike leaves. They are native to tropical Africa and islands ...

'' (formerly a subgenus of ''Antrophyum'') for three species placed in ''Scoliosorus'' but more distant from the type of that genus than ''Antrophyum''. This treatment also sank ''Anetium'' into ''Polytaenium'' and ''Monogramma'' into ''Haplopteris''. Since the name ''Monogramma'' has taxonomic priority over ''Haplopteris'', a proposal to reject ''Monogramma'' in favor of ''Haplopteris'' has been put forth to conserve the name and comparatively stable circumscription of ''Haplopteris''.

Genera

The following genera are recognized in the Pteridophyte Phylogeny Group classification of 2016 (PPG I): * ''

'' L. * ''

'' Underw. & Maxon * ''

'' (Benedict) Schuettp. * ''

Antrophyum ''Antrophyum'' is a genus of ferns in the family Pteridaceae.''Antrophyum''
USDA Germplasm Reso ...

'' Kaulf. * ''

'' C.Presl * ''

Hecistopteris ''Hecistopteris'' is a fern genus in the Vittarioideae subfamily of the Pteridaceae. Species The genus ''Hecistopteris'' contains the following species:Polytaenium ''Polytaenium'' is a genus of ferns in the subfamily Vittarioideae of the family Pteridaceae. Species are native to Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin ...

'' Desv. * ''

'' (Benedict) E.H.Crane * ''

'' Alston * ''

'' T.Moore * '' Vaginularia'' Fée * ''

'' Sm. The following phylogeny for the currently recognized genera of the subfamily was presented by Schuettpelz ''et al.'':

References

{{Taxonbar, from1=Q5873935, from2=Q19387804, from3=Q3735428 Pteridaceae Plant subfamilies