These authors contributed equally to this publicationThe 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.
Mountains of Anatolia are one of the main Mediterranean biodiversity hotspots and their richness in endemic species amounts for 30% of the flora. Two main factors may account for this high diversity: the complex orography and its role as refugia during past glaciations. We have investigated seven narrow endemics of Centaurea subsection Phalolepis from Anatolia by means of microsatellites and ecological niche modelling (ENM), in order to analyse genetic polymorphisms and getting insights into their speciation. Despite being narrow endemics, all the studied species show moderate to high SSR genetic diversity. Populations are genetically isolated, but exchange of genes probably occurred at glacial maxima (likely through the Anatolian mountain arches as suggested by the ENM). The lack of correlation between genetic clusters and (morpho) species is interpreted as a result of allopatric diversification on the basis of a shared gene pool. As suggested in a former study in Greece, post-glacial isolation in mountains would be the main driver of diversification in these plants; mountains of Anatolia would have acted as plant refugia, allowing the maintenance of high genetic diversity. Ancient gene flow between taxa that became sympatric during glaciations may also have contributed to the high levels of genetic diversity.
The Achillea millefolium aggregate is one of the most diverse polyploid complexes of the Northern hemisphere and has its western Eurasian boundary in the Iberian Peninsula. Four ploidy levels have been detected in A. millefolium, three of which have already been found in Iberia (diploid, hexaploid and octoploid), and a fourth (tetraploid) reported during the preparation of this paper. We collected a sample from 26 Iberian populations comprising all ploidy levels, and we used microsatellite markers analyzed as dominant in view of the high ploidy levels. Our goals were to quantify the genetic diversity of A. millefolium in the Iberian Peninsula, to elucidate its genetic structure, to investigate the differences in ploidy levels, and to analyse the dispersal of the species. The lack of spatial genetic structure recovered is linked to both high levels of gene flow between populations and to the fact that most genetic variability occurs within populations. This in turn suggests the existence of a huge panmictic yarrow population in the Iberian Peninsula. This is consistent with the assumption that recent colonization and rapid expansion occurred throughout this area. Likewise, the low levels of genetic variability recovered suggest that bottlenecks and/or founder events may have been involved in this process, and clonal reproduction may have played an important role in maintaining this genetic impoverishment. Indeed, the ecological and phenologic uniformity present in the A. millefolium agg. in Iberia compared to Eurasia and North America may be responsible for the low number of representatives of this complex of species present in the Iberian Peninsula. The low levels of genetic differentiation between ploidy levels recovered in our work suggest the absence of barriers between them.
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