Molecular evidence of reticulate evolution in the subgenus Plantago (Plantaginaceae).

Polyploidization is a frequent evolutionary event in plants that has a large influence on speciation and evolution of the genome. Molecular phylogenetic analyses of the taxonomically complex subgenus Plantago were conducted to elucidate intrasubgeneric phylogenetic relationships. A nuclear-encoding single-copy gene, SUC1 (1.0-1.8 kb), was sequenced in 24 taxa representing all five sections of the subgenus Plantago and two taxa from subgenus Coronopus as the outgroup. Fifteen known polyploids and one putative polyploid were sampled to examine polyploid origins and occurrence of reticulate evolution by cloning and sequence analysis of SUC1. Phylogenetic relationships were estimated using maximum parsimony, neighbor-joining, and Bayesian analyses. For the first time, our analysis provides a highly resolved phylogenetic tree. Subgenus Plantago formed a well-supported monophyletic clade. In contrast, alleles from polyploid species were scattered across the whole SUC1 phylogenetic tree, and some independent allopolyploids originated from hybridization between distant lineages. One reason for this taxonomic complexity can be attributed to reticulate evolution within the subgenus Plantago. Our results also suggest the possibility of two independent long-distance dispersals between the northern and southern hemispheres.