Dryopteridaceae

The Dryopteridaceae are a family of leptosporangiate ferns in the order Polypodiales. They are known colloquially as the wood ferns. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family is placed in the suborder Polypodiineae. Alternatively, it may be treated as the subfamily Dryopteridoideae of a very broadly defined family Polypodiaceae sensu lato.

Dryopteridaceae
Dryopteris carthusiana
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Division:	Polypodiophyta
Class:	Polypodiopsida
Order:	Polypodiales
Suborder:	Polypodiineae
Family:	Dryopteridaceae Herter (nom. cons.)
Subfamilies
Dryopteridoideae Elaphoglossoideae Polybotryoideae
Synonyms
Bolbitidaceae Ching 1978 Elaphoglossaceae Pichi-Sermolli 1968 Filicaceae de Jussieu 1789 nom. ill. Peranemataceae Ching 1940 non Buetschli 1884

The family contains about 1700 species and has a cosmopolitan distribution. Species may be terrestrial, epipetric, hemiepiphytic, or epiphytic. Many are cultivated as ornamental plants. The largest genera are Elaphoglossum (600+), Polystichum (260), Dryopteris (225), and Ctenitis (150). These four genera contain about 70% of the species. Dryopteridaceae diverged from the other families in eupolypods I about 100 million years ago. The fossil record appears to show the earliest members of the group emerging some 99 MYA during the Mid Cretaceous.

Description

The rhizomes are often stout, creeping, ascending, or erect, and sometimes scandent or climbing, with nonclathrate scales at apices. Fronds are usually monomorphic, less often dimorphic, or sometimes scaly or glandular, but less commonly hairy. Petioles have numerous round, vascular bundles arranged in a ring, or rarely as few as three; the adaxial bundles are largest. Veins are pinnate or forking, free to variously anastomosing; the areoles occur with or without included veinlets; sori are usually round, acrostichoid (covering the entire abaxial surface of the lamina) in a few lineages; usually indusiate, or sometimes exindusiate. Indusia, when present, are round-reniform or peltate. Sporangia have three-rowed, short to long stalks; spores are reniform, monolete, or winged.

Taxonomy

History

In 1990, Karl U. Kramer and coauthors defined the Dryopteridaceae broadly to include the present family, as well as the Woodsiaceae sensu lato, Onocleaceae, and most of Tectariaceae.Molecular phylogenetic studies found Kramer's version of the Dryopteridaceae to be polyphyletic, and it was split up by Smith and others in 2006. The inclusion of Didymochlaena, Hypodematium, and Leucostegia in the Dryopteridaceae is doubtful. If these three are excluded, then the family is strongly supported as monophyletic in cladistic analyses. Some authors have already treated these genera as outside of the Dryopteridaceae.

In 2007, a phylogenetic study of DNA sequences showed that Pleocnemia should be transferred from the Tectariaceae to the Dryopteridaceae. In 2010, in a paper on ferns, Arthrobotrya was resurrected from Teratophyllum. Later that year, Mickelia was described as a new genus.

Some species have been removed from the genus Oenotrichia because they do not belong there or even in the family Dennstaedtiaceae where Oenotrichia sensu stricto is placed. These species probably belong in the Dryopteridaceae, but have not yet been given a generic name.

In 2012, a phylogenetic study of Dryopteris and its relatives included Acrophorus, Acrorumohra, Diacalpe, Dryopsis, Nothoperanema, and Peranema within that genus. The Flora of China treatment of the family, published in 2013, used phylogenetic results to sink Lithostegia and Phanerophlebiopsis into Arachniodes.

The Dryopteridaceae Herter, under the classification system of Christenhusz and Chase (2014), were submerged as subfamily Dryopteridoideae Link, one of eight subfamilies constituting family Polypodiaceae. This family corresponds to the clade eupolypods I. The Pteridophyte Phylogeny Group classification of 2016 (PPG I) retained the family.

Phylogeny

The following cladogram for the suborder Polypodiineae (eupolypods I), based on the consensus cladogram in the Pteridophyte Phylogeny Group classification of 2016 (PPG I), shows a likely phylogenetic relationship between Dryopteridaceae and the other families of the clade.

Polypodiineae (eupolypods I)	Didymochlaenaceae Hypodematiaceae Dryopteridaceae Nephrolepidaceae Lomariopsidaceae Tectariaceae Oleandraceae Davalliaceae Polypodiaceae

Subdivision

Phylogeny of Dryopteridaceae

Elaphoglossoideae Parapolystichum Lastreopsis Rumohra Megalastrum Pleocnemia Bolbitis Lomagramma Arthrobotrya Teratophyllum Mickelia Elaphoglossum Polybotryoideae Stigmatopteris Trichoneuron Polystichopsis Olfersia Maxonia Cyclodium Polybotrya Dryopteridoideae Ctenitis Phanerophlebia Cyrtomium Polystichum Arachniodes Dryopteris

The PPG I classification divides the family into three subfamilies, listed below.

• Subfamily Polybotryoideae H.M.Liu & X.C.Zhang
  • Cyclodium C.Presl
  • Maxonia C.Chr.
  • Olfersia Raddi
  • Polybotrya Humb. & Bonpl. ex Willd.
  • Polystichopsis (J.Sm.) Holttum
  • Stigmatopteris C.Chr.
  • Trichoneuron Ching
• Subfamily Elaphoglossoideae (Pic.Serm.) Crabbe, Jermy & Mickel
  • Arthrobotrya J.Sm.
  • Bolbitis Schott
  • Elaphoglossum Schott ex J.Sm.
  • Lastreopsis Ching
  • Lomagramma J.Sm.
  • Megalastrum Holttum
  • Mickelia R.C.Moran, Labiak & Sundue
  • Parapolystichum (Keyserl.) Ching
  • Pleocnemia C.Presl
  • Rumohra Raddi
  • Teratophyllum Mett. ex Kuhn
• Subfamily Dryopteridoideae Link
  • Arachniodes Blume
  • Ctenitis (C.Chr.) C.Chr.
  • Cyrtomium C.Presl
  • Dryopteris Adans.
  • Phanerophlebia C.Presl
  • Polystichum Roth

Didymochlaena has been removed to Didymochlaenaceae, and Hypodematium and Leucostegia to Hypodematiaceae. Aenigmopteris has at times been suggested to belong to this family, on the grounds of its morphological similarity to Ctenitis, but molecular phylogeny has led to its submersion within Tectaria (Tectariaceae).Dryopolystichum has been placed in Lomariopsidaceae.

References

1. 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. S2CID 39980610.
2. Christenhusz, Maarten J.M. & Chase, Mark W. (2014). "Trends and concepts in fern classification". Annals of Botany. 113 (9): 571–594. doi:10.1093/aob/mct299. PMC 3936591. PMID 24532607.
3. Sue Olsen. 2007. Encyclopedia of Garden Ferns Timber Press: Portland, OR, USA. ISBN 978-0-88192-819-8
4. Smith et al., 2006 Archived February 26, 2008, at the Wayback Machine Alan R. Smith, Kathleen M. Pryer, Eric Schuettpelz, Petra Korall, Harald Schneider & Paul G. Wolf: "A classification for extant ferns," Taxon, 55(3): 705–731 (Aug 2006)
5. Eric Schuettpelz and Kathleen M. Pryer. 2009. "Evidence for a Cenozoic radiation of ferns in an angiosperm-dominated canopy". Proceedings of the National Academy of Sciences 106(27):11200-11205. doi:10.1073/pnas.0811136106
6. Long, Xiaoxuan; Peng, Yuan; Feng, Qi; Engel, Michael S.; Shi, Chao; Wang, Shuo (2023-09-01). "A new fossil fern of the Dryopteridaceae (Polypodiales) from the mid-Cretaceous Kachin amber". Palaeobiodiversity and Palaeoenvironments. 103 (3): 489–494. doi:10.1007/s12549-023-00572-4. ISSN 1867-1608.
7. Karl U. Kramer (with Richard E. Holttum, Robin C. Moran, and Alan R. Smith). 1990. "Dryopteridaceae". pages ??. 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
8. Alan R. Smith, Kathleen M. Pryer, Eric Schuettpelz, Petra Korall, Harald Schneider, and Paul G. Wolf. 2008. "Dryopteridaceae". pages ??. In: "Fern Classification". pages 417-467. 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
9. Christenhusz, Maarten J. M.; Zhang, Xian-Chun; Schneider, Harald (18 February 2011). "A linear sequence of extant families and genera of lycophytes and ferns" (PDF). Phytotaxa. 19: 7–54. doi:10.11646/phytotaxa.19.1.2. hdl:10138/28042. ISSN 1179-3163.
10. Hong-Mei Liu, Xian-Chun Zhang, Wei Wang, Yin-Long Qiu, and Zhi-Duan Chen. 2007. "Molecular Phylogeny of the Fern Family Dryopteridaceae inferred from Chloroplast rbcL and atpB genes". International Journal of Plant Sciences 168(9):1311-1323. doi:10.1086/521710
11. Robbin C. Moran, Paulo H. Labiak, and Michael Sundue. 2010. "Phylogeny and character evolution of the bolbitidoid ferns (Dryopteridaceae)". International Journal of Plant Sciences 171(5):547-559. doi:10.1086/652191
12. Robbin C. Moran, Paulo H. Labiak, and Michael Sundue. 2010. "Synopsis of Mickelia, a newly recognized genus of bolbitidoid ferns (Dryopteridaceae)". Brittonia 62(4):337-356.
13. Li-Bing Zhang, Liang Zhang, Shi-Yong Dong, and Atsushi Ebihara. 2012. "Molecular circumscription and major evolutionary lineages of the fern genus Dryopteris (Dryopteridaceae)". BMC Evolutionary Biology 12(1):180
14. He H, Wu SG, Xiang JY, Barrington DS (2013) "Arachniodes". In: Wu ZY, Raven PH, Hong DY (eds) Flora of China, vol 2–3.
15. Christenhusz & Chase 2014.
16. Nitta, Joel H.; Schuettpelz, Eric; Ramírez-Barahona, Santiago; Iwasaki, Wataru; et al. (2022). "An Open and Continuously Updated Fern Tree of Life". Frontiers in Plant Science. 13. doi:10.3389/fpls.2022.909768. PMC 9449725. PMID 36092417.
17. "Tree viewer: interactive visualization of FTOL". FTOL v1.3.0. 2022. Retrieved 12 December 2022.
18. Chen, Cheng-Wei; RothfelsE, Carl J.; Mustapeng, Andi Maryani A.; Gubilil, Markus; Karger, Dirk Nikolaus; Kessler, Michael; Huang, Yao-Moan (2018). "End of an enigma: Aenigmopteris belongs in Tectaria (Tectariaceae: Polypodiopsida)". Journal of Plant Research. 131 (1): 67–76. doi:10.1007/s10265-017-0966-9. PMID 28741041. S2CID 4573970.
19. Chen, Cheng-Wei; Sundue, Michael; Kuo, Li-Yaung; Teng, Wei-Chih; Huang, Yao-Moan (2017). "Phylogenetic analyses place the monotypic Dryopolystichum within Lomariopsidaceae". PhytoKeys (78): 83–107. doi:10.3897/phytokeys.78.12040. PMC 5543276. PMID 28781553.

Bibliography

• Smith, Alan R.; Pryer, Kathleen M.; Schuettpelz, Eric; Korall, Petra; Schneider, Harald; Wolf, Paul G. (1 January 2006). "A Classification for Extant Ferns" (PDF). Taxon. 55 (3): 705–731. doi:10.2307/25065646. JSTOR 25065646.

External links

• Phytotaxa