Gj. Gastony et Dr. Rollo, PHYLOGENY AND GENERIC CIRCUMSCRIPTIONS OF CHEILANTHOID FERNS (PTERIDACEAE, CHEILANTHOIDEAE) INFERRED FROM RBCL NUCLEOTIDE-SEQUENCES, American fern journal, 85(4), 1995, pp. 341-360
Nucleotide sequences of the chloroplast DNA gene rbcL were determined
for 25 species of cheilanthoid ferns (Pteridaceae subfamily Cheilantho
ideae). Together with GenBank sequences of an additional cheilanthoid
and a 14-species outgroup comprising Pteridaceae, Vittariaceae, and Co
niogramme japonica, these were analyzed cladistically by maximum parsi
mony to gain insights into cheilanthoid phylogeny and generic circumsc
riptions, which have long stymied pteridologists. Our analysis yielded
4 equally most parsimonious trees of 1570 steps. Two of these trees v
aried only in the relative positions of Onychium and Pteris in the glo
bal outgroup. The other two varied only in the relative positions of P
ellaea pringlei and P. rotundifolia in the ingroup. Results based on r
bcL sequences are concordant with those of recent studies of Argyrocho
sma, Hemionitis, and Bommeria that utilized a broad suite of character
s, suggesting that rbcl-based inferences in less meticulously studied
cheilanthoid groups also convey meaningful information. Among the insi
ghts into cheilanthoid phylogeny and generic circumscriptions offered
by rbcL sequences are the following. Llavea does not belong in subfami
ly Cheilanthoideae. Pellaea and Cheilanthes are polyphyletic. The rece
nt removal of 21 species from Notholaena to Cheilanthes and the segreg
ation of Argyrochosma are supported. The transfer of Hemionitis elegan
s to Bommeria is strongly supported, but the removal of the group of D
oryopteris concolor to Cheilanthes is not. Trachypteris is sister to D
oryopteris. The segregation of some small genera from Cheilanthes is s
upported, but these require further study through inclusion of additio
nal taxa. Bommeria is the most basal ingroup element in this analysis.