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Grammitis (dwarf polypody) is a genus of ferns in the family Polypodiaceae.[1] It had formerly been placed in the family Grammitidaceae, but this family is no longer recognized by most authors because phylogenetic analyses of DNA sequences have shown that it is embedded in Polypodiaceae.[2]

Finger Fern Grammitis billardierei - Mt Imlay.jpg
Grammitis billardierei
Scientific classification e
Kingdom: Plantae
Class: Polypodiopsida
Order: Polypodiales
(unranked): Eupolypods I
Family: Polypodiaceae
Subfamily: Grammitidoideae
Genus: Grammitis
O. Swartz
Type species
Grammitis marginella
(O. Swartz) O. Swartz

See text

The delimitation of Grammitis was drastically narrowed in the first decade of the 21st century. It now contains about 25 species. In 2003, a study of the distribution of grammitid ferns placed 11 species in the New World, 7 in Africa, and 4 in the Pacific.[3]

The genus Grammitis was established by Olof Swartz around 1801.[4] (sources vary on the exact date). The name is derived from Greek, gramma, grammatos meaning "a line or thread" and refers to the arrangement of the sori in some species.[5] The type species for Grammitis is Grammitis marginella.[6]

The only known fossil of a grammitid fern has been named Grammitis succinea, but it is not clear that it belongs to Grammitis as more recently defined. It was found in Oligocene amber from the Dominican Republic.[7]



In a treatment of Grammitidaceae in 1990, Barbara S. Parris defined Grammitis broadly, to include about 400 species.[8] At that time, she stated that "The treatment of Grammitis here as a large and diverse genus reflects our current lack of knowledge concerning relationships within the family". Other authors at that time circumscribed Grammitis more narrowly, to include about 200 species.[7]

Since about 2003, Grammitis has been understood as a genus of about 25 species with the other species to be eventually transferred to other genera, most of them new or resurrected.[3] Grammitis sensu strictissimo (about 25 species) is distinguished by its black, sclerified leaf margins.[6] It is sister to the monophyletic genus Cochlidium.[6]

In 2004, Grammitis was shown to be polyphyletic if broadly circumscribed.[9] Grammitis tenella had been shown to be closely related to Adenophorus in 2003,[10] and the new combination Adenophorus tenellus was published in 2008.[6] Several species formerly in Grammitis have been transferred to Oreogrammitis. These include Oreogrammitis clemensiae, Oreogrammitis hookeri, and others.[6][11] The genera Ctenopterella, Dasygrammitis, Radiogrammitis, Tomophyllum, and Xiphopterella were established in 2007 and some of these contain species that were formerly in Grammitis.[11] Lellingeria tomensis was moved out of Grammitis to Lellingeria in 2004.[12] Transfers to other genera have also been made. The classification of Grammitis basalis, Grammitis recondita, and others remains undecided.


The species list is incomplete and may contain synonyms. It was originally based on two very old sources.[13][14]

(=) Polypodium basale Maxon ex C.V.Morton 1962
(=) Polypodium billardierei C.Chr. 1906
(=) Grammitis haplophlebica (A.C.Sm.) Vareschi 1969
(=) Polypodium haplophlebicum A.C. Sm. 1931
(=) Polypodium bryophilum Maxon 1926
(=) Grammitis pseudomarginella (Bonap.) Copel. 1952
(=) Polypodium leptopodon C.H. Wright 1901
(=) Grammitis nigrolimbata (Spruce ex Jenman) Lellinger 1977
(=) Polypodium hessii Maxon 1915
(=) Polypodium limbatum (Fée) Maxon 1915
(=) Polypodium nigrolimbatum Spruce ex Jenman 1897 (nom. illeg.)
(=) Grammitis nigrolimbata Spruce ex Hook. 1862 (nom. inval., nom. nud.)
(=) Mecosorus marginellus (Sw.) Klotzsch 1847
(=) Polypodium marginellum Sw. 1788
(=) Polypodium marginelloides J.W. Moore 1933
(=) Polypodium melanoloma (Boiv.) Cordem. 1891
(=) Polypodium microglossum C.Chr. 1932
(=) Polypodium poeppigianum Mett. 1857
(=) Polypodium vaupelii Brause 1922

Transferred speciesEdit

(=) Grammitis tenella Kaulf. 1824
(=) Polypodium pseudogrammitis Gaudich. 1827


  1. ^ Eric Schuettpelz and Kathleen M. Pryer. 2008. "Fern phylogeny" pages 395-416. In: Tom A. Ranker and Christopher H. Haufler (editors). Biology and Evolution of Ferns and Lycophytes. Cambridge University Press. ISBN 978-0-521-87411-3
  2. ^ Harald Schneider, Hans-Peter Krier, Rosemary Wilson, and Alan R. Smith. 2006. "The Synammia Enigma: Evidence for a Temperate Lineage of Polygrammoid Ferns (Polypodiaceae, Polypodiidae) in Southern South America". Systematic Botany 31(1):31-41.
  3. ^ a b Barbara S. Parris. 2003. "The distribution of Grammitidaceae (Filicales) inside and outside Malesia". Telopea 10(2):451-466.
  4. ^ Olof Swartz. about 1801. "Genera et Species Filicum ···" In: Schrader's Journal für die Botanik. page 17. (see External links below).
  5. ^ Umberto Quattrocchi. 2000. CRC World Dictionary of Plant Names volume II. CRC Press: Boca Raton; New York; Washington,DC;, USA. London, UK. ISBN 978-0-8493-2676-9 (vol II). (see External links below).
  6. ^ a b c d e Tom A. Ranker. 2008. "A New Combination in Adenophorus (Polypodiaceae)". American Fern Journal 98(3):170-177.
  7. ^ a b David J. Mabberley. 2008. Mabberley's Plant-Book third edition (2008). Cambridge University Press: UK. ISBN 978-0-521-82071-4
  8. ^ Barbara S. Parris. 1990. "Grammitidaceae" pages 153-157. In: Klaus Kubitzki (general editor); Karl U. Kramer and Peter S. Green (volume editors) The Families and Genera of Vascular Plants volume I. Springer-Verlag: Berlin;Heidelberg, Germany. ISBN 978-0-387-51794-0
  9. ^ Tom A. Ranker, Alan R. Smith, Barbara S. Parris, Jennifer M.O. Geiger, Christopher H. Haufler, Shannon C.K. Straub, and Harald Schneider. 2004. "Phylogeny and evolution of grammitid ferns (Grammitidaceae): a case of rampant morphological homoplasy". Taxon 53(2):415-428.
  10. ^ Tom A. Ranker, Jennifer M.O. Geiger, S.C. Kennedy, Alan R. Smith, Christopher H. Haufler, and Barbara S. Parris. 2003. "Molecular phylogenetics and evolution of the endemic Hawaiian genus Adenophorus (Grammitidaceae)" Molecular Phylogenetics and Evolution 26(3):337-347.
  11. ^ a b Barbara S. Parris. 2007. "Five new genera and three new species of Grammitidaceae (Filicales) and the re-establishment of Oreogrammitis". Gardens' Bulletin. Singapore 58(2):233-274.
  12. ^ Barbara S. Parris. 2004. "New combinations in Acrosorus, Lellingeria, Prosaptia, and Themelium (Grammitidaceae: Filicales)" Kew Bulletin 59(2):223-225.
  13. ^ Edwin B. Copeland. 1956. Key to Subgenus Melanoloma page 253. In: "Grammitis" pages 93-278. In: Philippine Journal of Science 80(2).
  14. ^ L. Earl Bishop. 1977. "The American Species of Grammitis Sect. Grammitis". American Fern Journal 67(4):101-106.

External linksEdit