Phylogeny and evolution of the Arctium‐Cousinia complex (Compositae, Cardueae‐Carduinae)

Abstract

AbstractThe phylogeny and evolution of the Arctium‐Cousinia complex, including Arctium, Cousinia as one of the largest genera of Asteraceae, Hypacanthium and Schmalhausenia, is investigated. This group of genera has its highest diversity in the Irano‐Turanian region and the mountains of Central Asia. We generated ITS and rpS4‐trnT‐trnL sequences for altogether 138 species, including 129 (of ca. 600) species of Cousinia. As found in previous analyses, Cousinia is not monophyletic. Instead, Cousinia subgg. Cynaroides and Hypacanthodes with together ca. 30 species are more closely related to Arctium, Hypacanthium and Schmalhausenia (Arctioid clade) than to subg. Cousinia (Cousinioid clade). The Arctioid and Cousiniod clades are also supported by pollen morphology and chromosome number as reported earlier. In the Arctioid clade, the distribution of morphological characters important for generic delimitation, mainly leaf shape and armature and morphology of involucral bracts, are highly incongruent with phylogenetic relationships as implied by the molecular data. No taxonomic solution for this conflict can be offered, and the characters named are interpreted as homoplasious. Although phylogenetic resolution in the Cousinioid clade is poor, our ITS and rpS4‐trnT‐trnL sequences contain some phylogenetic information. For example, the six annual species of the Cousinioid clade fall into two groups. Poor phylogenetic resolution probably results from lack of characters and the high number of taxa in this species‐rich and comparatively young (ca. 8.7 mya) lineage. We hypothesize that speciation in the Cousinioid clade was mainly allopatric.