Polyphyly of the spring-parsleys (Cymopterus): molecular and morphological evidence suggests complex relationships among the perennial endemic genera of western North American Apiaceae

Cladistic analyses of DNA sequences from the nuclear rDNA internal transcribed spacer region and cpDNA rps16 intron and, for a subset of taxa, the cpDNA trnF-trnL-trnT locus were carried out to evaluate the monophyly of Cymopterus and to ascertain its phylogenetic placement among the other perennial genera of Apiaceae (Umbelliferae) subfamily Apioideae endemic to western North America. To elucidate patterns in the evolution of specific fruit characters and to evaluate their utility in circumscribing genera unambiguously, additional evidence was procured from cross-sections of mature fruits and the results of cladistic analysis of 25 morphological characters. Analyses of the partitioned data sets resulted in weakly supported and largely unresolved phylogenetic hypotheses, possibly due to the rapid radiation of the group, whereas the combined analysis of all molecular evidence resulted in a well-resolved phylogeny with higher bootstrap support. The traditionally used fruit characters of wing shape and composition and orientation of mericarp compression are highly variable. The results of these analyses reveal that Cymopterus and Lomatium, the two largest genera of western North American Apiaceae, are polyphyletic, and that their species are inextricably linked with those of other endemic perennial genera of the region (such as, Aletes, Musineon, Oreoxis, Pseudocymopterus, Pteryxia, and Tauschia), many of which are also not monophyletic. Prior emphasis on characters of the fruit in all systems of classification of the group has led to highly artificial assemblages of species. A complete reassessment of generic limits of all western endemic Apiaceae is required, as is further systematic study of this intractable group.