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Drynaria rigidula.jpg
Cultivated Drynaria rigidula in Florida
Scientific classification
Kingdom: Plantae
Division: Pteridophyta
Class: Pteridopsida
Order: Polypodiales
(unranked): Eupolypods I
Family: Polypodiaceae
Subfamily: Polypodioideae
B.K.Nayar 1970

sensu PPG I, 2016[1]

Polypodioideae is a subfamily belonging to the fern family Polypodiaceae. The subfamily name Polypodioideae has two recent uses. The first by Mabberley, 2008,[2] included a larger group of polygrams, (said to possess scales and/or hairs; never stellate), which included six tribes, one of which, Polypodieae, is considered to be a synonym of the more recent sense of Polypodioideae by Christenhusz et al., 2011,[3] in which Polypodiaceae contains five subfamilies, including the smaller Polypodioideae subfamily.


Mabberley, in 2008, defined this subfamily in a very broad sense, including all of Polypodiaceae except for the Platycerioideae (Platycerium and Pyrrosia) and the grammitid ferns, which he placed in Grammitidaceae. He distinguished this subfamily from the platycerioids by the absence of stellate hairs on the fronds. He subdivided it into six tribes, Drynarieae (fronds dimorphic; Aglaomorpha and Drynaria), Selligueeae (fronds monomorphic, with opaque scales on the stem; Selliguea), Lepisoreae (fronds monomorphic, with clathrate scales and thick exospore; Belvisia and Lepisorus), Microsoreae (fronds monomorphic, with clathrate scales and thin exospore), Polypodieae (Pecluma and Polypodium), and Loxogrammeae (lacking internal sclerenchyma above roots; Loxogramme).[2]

Christenhusz et al., in 2011, incorporated phylogenetic evidence to recircumscribe the family, combining the grammitids with Mabberley's Polypodieae (recognizing a number of segregate genera).[3] The classification of Christenhusz and Chase in 2014 vastly expanded the circumscription of Polypodiaceae; it redefined Polypodioideae to include all of what had generally been theretofore recognized as Polypodiaceae, including the grammitids. The authors recognized five tribes in this expanded circumscription: Loxogrammeae, Drynariae (including both drynarioid and selligueoid ferns), Platycerieae (platycerioids), Microsoreae (including Mabberley's Lepisoreae), and Polypodieae, the last corresponding to Polypodioideae in Christenhusz et al.[4]

The most recent circumscription of the subfamily, the PPG I classification, largely returns to that of Christenhusz et al., but excludes the grammitids as subfamily Grammitoideae, rendering the Polypodioideae paraphyletic.[1]


The following phylogram shows a likely relationship between Polypodioideae and the other subfamilies within Polypodiaceae, based on groups described by Schuettpelz & Pryer, 2008,[5] using the subfamily names of Christenhusz et al., 2011.[3]








"grammitid" ferns


The following grouping of Polypodioideae genera is based on Christenhusz et al., 2011.[3]

Non-grammitid genera[edit]

Grammitid genera[edit]

(includes family Grammitidaceae)

(=) Xiphopteris Kaulf. 1820
(=) Ctenopteris Blume ex Kunze[15]
  • Radiogrammitis Parris 2007 (Southeastern Asia, Pacific islands)
  • Scleroglossum Alderwerelt 1912 (Southeastern Asia, Pacific islands)
(=) Nematopteris Alderw.[15]
  • Stenogrammitis Labiak 2011 (Neotropics, Africa, Pacific islands)[16]
  • Terpsichore A.R.Sm. 1993 (Neotropics)
  • Themelium (T.Moore) Parris 1997 (China, Philippines, Thailand)
  • Tomophyllum (E.Fournier) Parris 2007 (India, Southeastern Asia, Australia)
  • Xiphopterella Parris 2007 (India, Southeastern Asia, China)
  • Zygophlebia L.E.Bishop 1989 (Neotropics, Madagascar)


  1. ^ a b The Pteridophyte Phylogeny Group (November 2016). "A community-derived classification for extant lycophytes and ferns". Journal of Systematics and Evolution. 54 (6): 563–603. doi:10.1111/jse.12229. 
  2. ^ a b D. J. Mabberley (2008). Mabberley's plant-book: a portable dictionary of plants, their classification and uses. Cambridge University Press. p. 690. ISBN 978-0-521-82071-4. 
  3. ^ a b c d Christenhusz et al. 2011 Maarten J. M. Christenhusz, Xian-Chun Zhang & Herald Schneider: "A linear sequence of extant families and genera of lycophytes and ferns." Phytotaxa 19: 7-54. (18 Feb. 2011)
  4. ^ Christenhusz, Maarten J. M.; Chase, Mark W. (13 February 2014). "Trends and concepts in fern classification". Annals of Botany. 113 (4): 571–594. doi:10.1093/aob/mct299. 
  5. ^ Schuettpelz & Pryer, 2008 Eric Schuettpelz & Kathleen M. Pryer: "Ch. 15. Fern phylogeny" in Biology and Evolution of Ferns and Lycophytes, ed. Tom A. Ranker and Christopher H. Haufler. Cambridge University Press (2008)
  6. ^ Ranker et al. 2003 T.A. Ranker, J.M.O. Geiger, S.C. Kennedy, A.R. Smith, C.H. Haufler, and B.S. Parris: "Molecular phylogenetics and evolution of the endemic Hawaiian genus Adenophorus (Grammitidaceae)." Molecular Phylogenetics and Evolution 26 (2003) 337-47.
  7. ^ Kessler et al. 2011 Michael Kessler, Ana Laura Moguel Velázquez, Michael Sundue, Paulo H. Labiak: "Alansmia, a new genus of grammitid ferns (Polypodiaceae) segregated from Terpsichore." Brittonia 63(2): 233–244 (June 2011)
  8. ^ [ Parris 2013] Barbara S. Parris: "Archigrammitis, a new genus of grammitid fern (Polypodiaceae) from Malesia and Polynesia." Fern Gazette 19(4): 135–138 (2013)
  9. ^ Sundue, 2010 Michael A. Sundue: "A monograph of Ascogrammitis, a new genus of grammitid ferns (Polypodiaceae)." Brittonia 62(4): 357–399 (December 2010)
  10. ^ Sundue et al. 2012 Michael A. Sundue, Paulo H. Labiak, Julián Mostacero, Alan R. Smith: "Galactodenia, a new genus of grammitid ferns segregated from Terpsichore (Polypodiaceae)." Systematic Botany 37(2): 339–346 (April–June 2012)
  11. ^ Labiak et al. 2010 Paulo H. Labiak, Germinal Rouhan & Michael Sundue: "Phylogeny and taxonomy of Leucotrichum (Polypodiaceae): A new genus of grammitid ferns from the Neotropics." Taxon 59(3): 911–921 (June 2010)
  12. ^ Hirai et al. 2011 Regina Y. Hirai, Germinal Rouhan, Paulo H. Labiak, Tom A. Ranker & Jefferson Prado: "Moranopteris: A new Neotropical genus of grammitid ferns (Polypodiaceae) segregated from Asian Micropolypodium." Taxon 60(4): 1123–1137 (August 2011)
  13. ^ Sundue, 2012 Michael A. Sundue: "Mycopteris: a new neotropical genus of grammitid ferns (Polypodiaceae)." Brittonia 66(2): 174–185 (June 2014)
  14. ^ Perrie & Parris 2012 Leon M. Perrie & Barbara S. Parris: "Chloroplast DNA sequences indicate the grammitid ferns (Polypodiaceae) in New Zealand belong to a single clade, Notogrammitis gen. nov." New Zealand Journal of Botany 50(4): 457–475 (December 2012)
  15. ^ a b Christenhusz & Schneider, 2011 Maarten J. M. Christenhusz & Harald Schneider: "Corrections to Phytotaxa 19: Linear sequence of lycophytes and ferns," Phytotaxa (14 September 2011)
  16. ^ Labiak 2011 Paulo H. Labiak: "Stenogrammitis, a new genus of grammitid ferns segregated from Lellingeria (Polypodiaceae)." Brittonia 63(1): 141–145 (March 2011)