Unique expression of a sporophytic character on the gametophytes of notholaenid ferns (Pteridaceae).

Journal Article (Journal Article)

Premise of the study

Not all ferns grow in moist, shaded habitats; some lineages thrive in exposed, seasonally dry environments. Notholaenids are a clade of xeric-adapted ferns commonly characterized by the presence of a waxy exudate, called farina, on the undersides of their leaves. Although some other lineages of cheilanthoid ferns also have farinose sporophytes, previous studies suggested that notholaenids are unique in also producing farina on their gametophytes. For this reason, consistent farina expression across life cycle phases has been proposed as a potential synapomorphy for the genus Notholaena. Recent phylogenetic studies have shown two species with nonfarinose sporophytes to be nested within Notholaena, with a third nonfarinose species well supported as sister to all other notholaenids. This finding raises the question: are the gametophytes of these three species farinose like those of their close relatives, or are they glabrous, consistent with their sporophytes?


We sowed spores of a diversity of cheilanthoid ferns onto culture media to observe and document whether their gametophytes produced farina. To place these species within a phylogenetic context, we extracted genomic DNA, then amplified and sequenced three plastid loci. The aligned data were analyzed using maximum likelihood to generate a phylogenetic tree.

Key results

Here we show that notholaenids lacking sporophytic farina also lack farina in the gametophytic phase, and notholaenids with sporophytic farina always display gametophytic farina (with a single exception). Outgroup taxa never displayed gametophytic farina, regardless of whether they displayed farina on their sporophytes.


Notholaenids are unique among ferns in consistently expressing farina across both phases of the life cycle.

Full Text

Duke Authors

Cited Authors

  • Johnson, AK; Rothfels, CJ; Windham, MD; Pryer, KM

Published Date

  • June 2012

Published In

Volume / Issue

  • 99 / 6

Start / End Page

  • 1118 - 1124

PubMed ID

  • 22542903

Electronic International Standard Serial Number (EISSN)

  • 1537-2197

International Standard Serial Number (ISSN)

  • 0002-9122

Digital Object Identifier (DOI)

  • 10.3732/ajb.1200049


  • eng