Onocleaceae

Onocleaceae
Scientific classification
Kingdom:	Plantae
Division:	Pteridophyta
Class:	Polypodiopsida /  Pteridopsida (disputed)
Order:	Polypodiales
(unranked):	Eupolypods II
Family:	Onocleaceae
Genera
Onoclea Onocleopsis Matteuccia Pentarhizidium

Onocleaceae is a small family of terrestrial ferns in the Polypodiales order of the Polypodiopsida class.^[1] The family may contain from one to four genera of onocleoids, consisting of five species largely in north temperate climes. The four genera: Matteuccia, Onoclea, Onocleopsis, and Pentarhizidium may be included under the single genus Onoclea.^[2]

Members of the family have the following characteristics, being distinguished by having strongly dimorphic fronds, with the fertile fronds different from the sterile fronds. The rhizomes are long- to short-creeping to ascending, and sometimes stoloniferous (Matteuccia and Onocleopsis). The leaves are strongly dimorphic and the petioles have two vascular bundles uniting distally into a gutter-shape. The blades are pinnatifid or pinnate-pinnatifid. The veins are free or anastomosing, lacking included veinlets. The spores are reniform, brownish to green. The sori are enclosed (sometimes tightly) by reflexed laminar margins, also with membranous, often fugacious true indusia.^[1]

Formerly, the two species in the genus Pentarhizidium were considered to be members of Matteuccia, but genetic analysis has determined that they compose a basal sister clade to the rest of the family. This family has been determined by genetic analysis to be closely allied to the Blechnaceae, within the clade of families sometimes known as Blechnales (which includes the athyrioid ferns and asplenioid ferns as well) (this clade is often treated as part of the order Polypodiales however). Matteuccia struthiopteris was previously classified under the Dryopteridaceae, and still is by the USDA. ^[3]

Species

Five species, in four genera ^[4]

• Matteuccia struthiopteris (ostrich fern) - wide distribution including North America, Europe, and Asia
• Onoclea sensibilis (sensitive fern) - North America and eastern Asia, with the American and Asian populations being recognized as distinct varieties
• Onocleopsis hintonii - found in a restricted range of wet mountain canyons in southern Mexico and Guatemala; some botanists (especially Masahiro Kato) consider this to be a species of Matteuccia
• Pentarhizidium
  • Pentarhizidium orientale - eastern Asia as far south as the Himalayas
  • Pentarhizidium intermedia - China to India

Phylogenic relationships

The following phylogram shows a likely relationship between the Onocleaceae and the other families within eupolypods II, based on Lehtonen, 2011,^[5] and Rothfels & al., 2012.^[6]

eupolypods II	Cystopteridaceae Rhachidosoraceae Diplaziopsidaceae Aspleniaceae Hemidictyaceae Thelypteridaceae Woodsiaceae Onocleaceae Blechnaceae Athyriaceae

Recent taxonomic treatment has suggested that the four Onocleacae genera are so closely related that they should be included under the single genus Onoclea.^[2] A possible phylogenic relationship between Onocleaceae species (line lengths are not significant):

Onocleaceae	Matteuccia struthiopteris (or Onoclea struthiopteris) Onocleopsis hintonii (or Onoclea hintonii) Onoclea sensibilis var interrupta Onoclea sensibilis var sensibilis Pentarhizidium orientale (or Onoclea orientalis) Pentarhizidium intermedium (or Onoclea intermedia)

References

1. Alan R. Smith, Kathleen M. Pryer, Eric Schuettpelz, Petra Korall, Harald Schneider & Paul G. Wolf (2006). "A classification for extant ferns" (PDF). Taxon 55 (3): 705–731. doi:10.2307/25065646.
2. Maarten J. M. Christenhusz, Xian-Chun Zhang & Harald Schneider (2011). "A linear sequence of extant families and genera of lycophytes and ferns" (PDF). Phytotaxa 19: 7–54.
3. US Department of Agriculture. Natural Resources Conservation Service. Matteuccia struthiopteris
4. GJ Gastony and MC Ungerer (1997), "Molecular systematics and a revised taxonomy of the onocleoid ferns (Dryopteridaceae: Onocleeae)", American Journal of Botany (Botanical Society of America) 84 (6): 840–849, doi:10.2307/2445820, JSTOR 2445820
5. Samuli Lehtonen (2011). "Towards Resolving the Complete Fern Tree of Life" (PDF). PLoS ONE 6 (10): e24851. doi:10.1371/journal.pone.0024851. PMC 3192703. PMID 22022365.
6. Carl J. Rothfels, Anders Larsson, Li-Yaung Kuo, Petra Korall, Wen- Liang Chiou, Kathleen M. Pryer (2012). "Overcoming Deep Roots, Fast Rates, and Short Internodes to Resolve the Ancient Rapid Radiation of Eupolypod II Ferns". Systematic Biology 61 (1): 70. doi:10.1093/sysbio/sys001. PMID 22223449.