Phylogenomics, floral evolution, and biogeography of Lithospermum L. (Boraginaceae).

Lithospermum (Boraginaceae), a geographically cosmopolitan medium-sized genus, includes diverse floral morphology, with variation in corolla size and shape and in breeding system. Over the past decade, multiple studies have examined the evolutionary history of Lithospermum, with most utilizing DNA regions from the plastid genome and/or the nuclear ribosomal internal transcribed spacer. These studies have, in general, not resulted in well-resolved and well-supported phylogenies. In the present study, 298 nuclear DNA regions, amplified via target sequence capture, were utilized for phylogenetic reconstruction for Lithospermum and relatives in Boraginaceae, and patterns of floral evolution, species diversification, and biogeography were examined. Based on multiple phylogenetic methods, Lithospermum is resolved as monophyletic, and the New World species of the genus are also monophyletic. While minimal phylogenetic incongruence is resolved within the nuclear genome, incongruence between the nuclear and plastid genomes is recovered. This is likely due to incomplete lineage sorting during early diversification of the genus in the Americas approximately 7.8 million years ago. At least four shifts to longer corollas are identified throughout Lithospermum, and this may be due to selection for hummingbird-pollinated flowers, particularly for species in Mexico and the southwestern United States. In the New World, one clade of species of the genus diversified primarily across the United States and Canada, and another radiated throughout the mountains of Mexico.