Flora of North America species comparison

Cystopteris is a taxonomically difficult genus at the species level. Especially troublesome is the worldwide and polymorphic species C. fragilis sensu lato. To maintain it as a single species with several varieties would be easiest (and least controversial). This approach, however, may not accurately reflect true evolutionary history.

Although Cystopteris species are found in temperate climates worldwide at tetraploid to octaploid ploidy levels, extant diploid species are concentrated in North America. The diploid species are relatively distinct from one another and are the progenitors of numerous allopolyploid derivatives (see reticulogram). In addition, an extinct (or undiscovered) diploid may have been involved in the origin of some polyploids (shown as "C. hemifragilis" on the reticulogram).

Considerable overlap exists among the leaf morphologies in the species of Cystopteris, even among the diploid taxa. Consequently, the key requires observation of subtle and sometimes overlapping characteristics.

Several general recommendations can be made for identifying Cystopteris. (1) Field workers should be aware that whenever Cystopteris species occur together, hybridization is likely; hybrids usually have shriveled and malformed spores. (2) Species of Cystopteris frequently occur as highly reduced plants, especially in stressful habitats such as high elevations, high latitudes, and cold and/or dry climates. Such stunted plants can be fertile, but leaf and stem characters required to distinguish species can be obscured. (3) Because of the importance of examining stem and spore features in distinguishing species, collectors should always attempt to obtain complete, fertile specimens.

Species ca. 20 (9 in the flora).

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Cystopteris fragilis is most often confused with C. tenuis in the east and C. reevesiana in the southwest. Habitat and geography, as well as the morphologic features discussed in the key, usually serve to separate these taxa. For instance, C. fragilis is more likely to be found on cliffs whereas the other species prefer boulders and soil (C. fragilis occurs at higher elevations and/or latitudes than the other species). These distinctions can be confounded when C. fragilis forms hybrids with sympatric species. Sterile pentaploid plants have been discovered where C. fragilis overlaps with C. laurentiana, tetraploid hybrids are likely where C. fragilis occurs with C. tenuis, and triploids may form where C. fragilis is found with C. reevesiana. Even after segregating relatively distinct elements such as Cystopteris protrusa, C. reevesiana, and C. tenuis, and identifying sterile hybrids, C. fragilis still remains a polymorphic and complex taxon that probably contains a number of natural, cryptic evolutionary units. For example, morphologically distinct hexaploid cytotypes have been reported (C. H. Haufler and M. D. Windham 1991). These occur as isolated and disjunct populations in Ontario, Alaska, Arizona, Colorado, Montana, and Wyoming. Isozymic profiles of each of these populations indicate that the hexaploids are polyphyletic and should not be accorded species status.

The presence of verrucate spores (as opposed to the normal spiny spores) has been used to circumscribe Cystopteris dickieana. Although genetic analyses have not been undertaken, we think the verrucate spore is probably a recessive feature controlled by one or a few genes. While present at low frequency in much of the range of C. fragilis, verrucate spores are particularly prominent in the Great Plains. Perhaps in this region the genetic combinations specifying verrucate spores have been fixed. Following R. F. Blasdell (1963), C. dickieana is also considered here to be conspecific with C. fragilis because (1) early stages in the development of spiny spores can appear verrucate (A. C. Jermy and L. Harper 1971), (2) the hexaploid cytotypes discussed above always have verrucate spores, regardless of their parentage, (3) individuals with verrucate spores can be found in populations that are otherwise uniformly spiny-spored, and (4) individuals and populations that have verrucate spores are not otherwise (morphologically, ecologically, or genetically) distinct from those that have spiny spores.

Especially in the western portion of its North American range (British Columbia, Washington, Montana, Idaho, Oregon, California), Cystopteris fragilis appears to be developing morphologically and ecologically distinctive variants. Hybrid individuals with aborted spores have been discovered, and plants from these areas increasingly tend to grow on both soil and rock and to have slightly different morphologies on the two substrates. These variants intergrade, however, and are not sufficiently distinct to warrant species status. This polymorphic polyploid is probably actively speciating at the tetraploid level, perhaps through gene silencing (C. R. Werth and M. D. Windham 1991).

(Discussion copyrighted by Flora of North America; reprinted with permission.)