Pteridaceae
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Pteridaceae is 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 t ...
s 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 ...
, 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 recognized as separate families: the adiantoid, cheilanthoid, pteridoid, and hemionitidoid ferns. Relationships among these groups remain unclear, and although some recent genetic analyses of the Pteridales suggest that neither the family Pteridaceae nor the major groups within it are all
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
, as yet these analyses are insufficiently comprehensive and robust to provide good support for a revision of the order at the family level.


Description

Members of Pteridaceae have creeping or erect
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 hori ...
s. The
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
are almost always compound and have linear sori that are typically on the margins of the
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
and lack a true
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 ...
, typically being protected by a false indusium formed from the reflexed margin of the leaf.


Taxonomy


Traditional groups

Curtis's botanical magazine, '' Argyrochosma nivea'' var. ''tenera'' As traditionally defined, the groups within Pteridaceae are as follows: * Adiantoid ferns (tribe ''Adianteae'' ); epipetric, terrestrial 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 ...
in moist habitats, rachis often dichotomously branching; sori relatively small and discrete with
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 life cyc ...
born on the false indusium rather than the leaf blade proper; only one 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 ...
'' L. – maidenhair ferns * Cheilanthoid ferns; primarily epipetric in semiarid habitats; leaves mostly with well-developed scales or
trichome Trichomes (); ) are fine outgrowths or appendages on plants, algae, lichens, and certain protists. They are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae. A covering of any kind of hair on a pla ...
s, often bipinnate or otherwise highly compound; sporangia mostly born in marginal sori with false indusia that are +/- continuous around the leaf margins; several genera, including: ** '' Argyrochosma'' (J.Sm.) Windham – false cloak ferns ** '' Aspidotis'' (Nutt. ex Hook.) Copel. – lace ferns ** ''
Astrolepis ''Astrolepis'' is a small genus of ferns in the family Pteridaceae. It was formed in 1992 from species previously placed in ''Cheilanthes'' and ''Notholaena''. The name is derived from the Greek words ἄστρον (), meaning "star," and λεπ ...
'' D.M.Benham & Windham – cloak ferns ** '' Cheilanthes'' Sw. – lip ferns ** '' Myriopteris'' Fée – lace lip ferns ** ''
Notholaena ''Notholaena'' (from Ancient Greek νόθο(ς) + χλαῖνα), cloak fern, is a genus of ferns in the Cheilanthoideae subfamily of the Pteridaceae. Ferns of this genus are mostly epipetric (growing on rock) or occurring in coarse, gravelly soi ...
'' R.Br. – cloak ferns ** '' Pellaea'' Link – cliff brakes * Pteridoid ferns (tribe ''Pterideae'' ); terrestrial and epipetric in moist habitats; leaves mostly without prominent scales or trichomes, most often
pinnate 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 ...
but sometimes more compound; sporangia born in marginal sori with false indusia that are +/- continuous around the leaf margins; several genera, including: ** ''
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 ...
'' L. – brakes ** '' Onychium'' Kaulf. * Parkerioid ferns (tribe ''Parkerieae'' ); aquatic in swamps and/or mangroves, including: ** '' Acrostichum'' L. – leather ferns ** '' Ceratopteris'' Brongn. * Hemionitidoid ferns; terrestrial, epipetric or epiphytic in moist or semiarid habitats; leaves simple, pinnate, or more compound; sporangia born in linear non-marginal, exindusiate sori or sometimes in marginal sori; several genera, including: ** '' Anogramma'' Link ** '' Cryptogramma'' R.Br. – rock brakes ** ''
Eriosorus ''Jamesonia'' is a genus of ferns in the subfamily Pteridoideae of the family Pteridaceae. It now includes the formerly separate genus ''Eriosorus''. Description Species of ''Jamesonia'' are terrestrial or grow on rocks. They vary considerably ...
'' Fée ** ''
Hemionitis ''Hemionitis'' is a genus of ferns in the subfamily Cheilanthoideae of the family Pteridaceae. Its circumscription varies greatly in different systems of fern classification. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), it ...
'' L. ** ''
Jamesonia ''Jamesonia'' is a genus of ferns in the subfamily Pteridoideae of the family Pteridaceae. It now includes the formerly separate genus ''Eriosorus''. Description Species of ''Jamesonia'' are terrestrial or grow on rocks. They vary considerably ...
'' Hook. & Grev. ** ''
Pityrogramma ''Pityrogramma'', the silverback ferns, or goldback ferns, is a fern genus in the subfamily Pteridoideae of the family Pteridaceae Pteridaceae is a family of ferns in the order Polypodiales, including some 1150 known species in ca 45 genera ...
'' Link – gold ferns * Vittarioid ferns (tribe ''Vittarieae'' ); primarily epiphytic in tropical regions and all have simple leaves with sori that follow the veins and lack true indusia: ** '' Anetium'' ** '' Antrophyum'' ** '' Hecistopteris'' ** '' Monogramma'' ** '' Vittaria'' – Shoestring fern


Subfamilies

Based on phylogenetic research, Christenhusz ''et al.'' (2011) divided the Pteridaceae genera into five subfamilies. These roughly correspond with the groups listed above, with the main difference being that ''adiantoid'' and ''vittarioid'' ferns are combined under the Vittarioideae subfamily name. The approach was followed 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 relatio ...
classification of 2016 (PPG I). * Cryptogrammoideae :Genera: ''
Coniogramme ''Coniogramme'' is one of three genera in the subfamily Cryptogrammoideae of the fern family Pteridaceae. A cultivated species, '' Coniogramme pilosa'', is known as "bamboo fern." Species *'' Coniogramme affinis'' ( C. Presl) Wall. ex Hieron ...
'', '' Cryptogramma'', '' Llavea'' * Parkerioideae :Genera: '' Acrostichum'', '' Ceratopteris'' * Pteridoideae :Genera: '' Actiniopteris'', '' Anogramma'', '' Austrogramme'', '' Cerosora'', '' Cosentinia'', '' Gastoniella'', ''
Jamesonia ''Jamesonia'' is a genus of ferns in the subfamily Pteridoideae of the family Pteridaceae. It now includes the formerly separate genus ''Eriosorus''. Description Species of ''Jamesonia'' are terrestrial or grow on rocks. They vary considerably ...
'' (incl. ''Eriosorus'' and ''Nephopteris''), '' Onychium'', ''
Pityrogramma ''Pityrogramma'', the silverback ferns, or goldback ferns, is a fern genus in the subfamily Pteridoideae of the family Pteridaceae Pteridaceae is a family of ferns in the order Polypodiales, including some 1150 known species in ca 45 genera ...
'', ''
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 ...
'' (incl. ''Neurocallis'' & ''Platyzoma''), '' Pterozonium'', '' Syngramma'', '' Taenitis'', '' Tryonia'' * Cheilanthoideae :Genera: ''
Adiantopsis ''Adiantopsis'' is a genus of ferns in the subfamily Cheilanthoideae of the 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), d ...
'', ''
Aleuritopteris ''Aleuritopteris'' is a genus of ferns in the Cheilanthoideae subfamily of the Pteridaceae. As with some other genera of the Cheilanthoideae, molecular phylogenetic studies have suggested that it is not monophyletic, and so may need to be circ ...
'', '' Allosorus'', '' Argyrochosma'', '' Aspidotis'', ''
Astrolepis ''Astrolepis'' is a small genus of ferns in the family Pteridaceae. It was formed in 1992 from species previously placed in ''Cheilanthes'' and ''Notholaena''. The name is derived from the Greek words ἄστρον (), meaning "star," and λεπ ...
'', '' Bommeria'', ''
Calciphilopteris ''Calciphilopteris'' is a genus of ferns in the family Pteridaceae. It is native to India and China, southward to Australia.Jovita C. Yesilyurt and Harald Schneider. 2010. "The new fern genus ''Calciphilopteris'' (Pteridaceae)". ''Phytotaxa'' 7 ...
'', '' Cheilanthes'', ''
Cheiloplecton ''Cheiloplecton'' is a genus of ferns in the subfamily Cheilanthoideae of the family Pteridaceae with a single species ''Cheiloplecton rigidum'', native to Mexico Mexico (Spanish: México), officially the United Mexican States, is a count ...
'', ''
Doryopteris ''Doryopteris'' is a genus of ferns in the subfamily Cheilanthoideae of the family Pteridaceae. Species The circumscription Circumscription may refer to: *Circumscribed circle * Circumscription (logic) *Circumscription (taxonomy) *Circumscrip ...
'', ''
Gaga Gaga ( he, גע גע literally 'touch touch') (also: ga-ga, gaga ball, or ga-ga ball) is a variant of dodgeball that is played in a gaga "pit". The game combines dodging, striking, running, and jumping, with the objective of being the last perso ...
'', ''
Hemionitis ''Hemionitis'' is a genus of ferns in the subfamily Cheilanthoideae of the family Pteridaceae. Its circumscription varies greatly in different systems of fern classification. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), it ...
'', '' Lytoneuron'', ''
Mildella ''Mildella'' is a genus of ferns in the subfamily Cheilanthoideae of the family Pteridaceae. Species are native to Mexico, Central America and the Caribbean. Species The Pteridophyte Phylogeny Group classification of 2016 (PPG I) placed seven s ...
'', '' Myriopteris'', ''
Notholaena ''Notholaena'' (from Ancient Greek νόθο(ς) + χλαῖνα), cloak fern, is a genus of ferns in the Cheilanthoideae subfamily of the Pteridaceae. Ferns of this genus are mostly epipetric (growing on rock) or occurring in coarse, gravelly soi ...
'', ''
Ormopteris ''Ormopteris'' is a genus of ferns in the subfamily Cheilanthoideae of the family Pteridaceae. Species are native to Brazil and Venezuela Venezuela (; ), officially the Bolivarian Republic of Venezuela ( es, link=no, República Bolivaria ...
'', '' Paragymnopteris'', '' Parahemionitis'', '' Pellaea'', '' Pentagramma'', '' Trachypteris'' *
Vittarioideae Vittarioideae is a subfamily of the fern family Pteridaceae, in the order Polypodiales. The subfamily includes the previous families Adiantaceae (adiantoids or maidenhair ferns) and Vittariaceae (vittarioids or shoestring ferns). Description The ...
:Genera: ''
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 ...
'', ''
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 ...
'', '' Antrophyopsis'', '' Antrophyum'', '' Haplopteris'', '' Hecistopteris'', '' Polytaenium'', '' Radiovittaria'', '' Rheopteris'', '' Scoliosorus'', '' Vaginularia'', '' Vittaria''


Phylogenic relationships

Smith et al. (2006) carried out the first higher-level
pteridophyte A pteridophyte is a vascular plant (with xylem and phloem) that disperses spores. Because pteridophytes produce neither flowers nor seeds, they are sometimes referred to as "cryptogams", meaning that their means of reproduction is hidden. Ferns, ...
classification published in the
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 ...
era. Smith referred to the ferns as monilophytes, dividing them into four groups. The vast majority of ferns were placed in the
Polypodiopsida 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 ...
. In 2016, the Pteridophyte Phylogeny Group divided 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 ...
into six suborders. Pteridaceae is the sole family in suborder Pteridiineae, with 52
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
. The suborder has the same
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 ...
as Smith et al. used for the family. The
phylogenetic 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 these six suborders is shown in this
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
: The oldest fossil of the family is '' Heinrichsia'' from the early Late Cretaceous (
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
) aged
Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. The ...
of
Myanmar Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John C. Wells, Joh ...
, which cannot be assigned to modern grouping of the family. Molecular clock evidence suggests a diversification of the family during the Late Cretaceous. Phylogeny of Pteridaceae.


Distribution and habitat

Mostly terrestrial or epipetric (growing on rock).


References


Bibliography

* * * * * * * * * * * * , in
Eric Schuettpelz. The evolution and diversification of epiphytic ferns. PhD Thesis Duke University 2007


* ttps://web.archive.org/web/20160304060354/http://www.anbg.gov.au/fern/taxa/classification.html Australian National Botanic Gardens. A classification of the ferns and their allies


External links


Tree of Life: Pteridaceae
{{Taxonbar, from=Q1141315 Fern families Polypodiales