Vittarioideae is a subfamily of the

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 t ...

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 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 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, is a genus of about 250 species of ferns in the subfamily Vittarioideae of the family Pteridaceae, though some researchers place it in its own family, Adiantaceae. The genus name comes from Greek, meaning "un ...

'', 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 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 ...

in tropical regions and all have simple leaves with sori that follow the veins and lack true

indusia 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 Gr ...

; the sori are most often marginal with a false indusium formed from the reflexed leaf margin. The family also includes a species, '' Vittaria appalachiana'', that is highly unusual in that the

sporophyte A sporophyte () is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase. Life cycle The sporophyte develops from the zygote pr ...

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

photosynthetic Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in c ...

gametophyte A gametophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has on ...

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, the circumscribed circle or circumcircle of a polygon is a circle that passes through all the vertices of the polygon. The center of this circle is called the circumcenter and its radius is called the circumradius. Not every polyg ...

, making that genus paraphyletic. In 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), 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 ( cs, Karel Bořivoj 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).

in 1836, who erected the

tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English language, English is in the discipline of anthropology. This definition is contested, in p ...

Vittariaceae to contain the genera ''Vittaria'' and '' Prosaptia'', the latter now included in the

grammitid Grammitidoideae is a subfamily of the fern family Polypodiaceae, whose members are informally known as grammitids. It comprises a clade of about 750 species. They are distributed over higher elevations in both the Old and New World. This group w ...

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 an ...

'', 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'', ''

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

'', '' Hecistopteris'', '' Monogramma'', '' Polytaenium'', and ''

''. He described '' Radiovittaria'' as a subgenus of ''Vittaria'', subsumed '' Scoliosorus'' within '' Polytaenium'' as doubtfully worthy of subgeneric rank, while '' Rheopteris'' 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 * Observational study * Research * Study skills, abilities and approaches applied to learning Other * Study (art), a drawing or series of drawi ...

of the vittarioids was based on the

chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...

gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...

rbcL Ribulose-1,5-bisphosphate carboxylase-oxygenase, commonly known by the abbreviations RuBisCo, rubisco, RuBPCase, or RuBPco, is an enzyme () involved in the first major step of carbon fixation, a process by which atmospheric carbon dioxide is con ...

. In this study, it was found that the

type species In 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 species that contains the biological type specimen ...

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'' (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 *'' Antrophyopsis'' (Benedict) Schuettp. *''

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

'' Kaulf. *''

'' C.Presl *'' Hecistopteris'' J.Sm. *'' Polytaenium'' Desv. *'' Radiovittaria'' (Benedict) E.H.Crane *'' Rheopteris'' Alston *'' Scoliosorus'' 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