Phylogenetics of the Paleartic model grass Brachypodium sylvaticum uncovers two divergent oriental and occidental micro-taxa lineages

Brachypodium sylvaticum has been selected as a model for perennial grasses, and considerable genomic resources have been generated and a reference genome and several resequenced pangenome accessions are available for this species. Despite these genomic advances, the evolution and systematics of diploid B. sylvaticum s. l. is almost unknown. The B. sylvaticum complex is formed by up to seven taxonomically close micro-taxa which differentiate from typical B. sylvaticum s. s. based on a few morphological features. Moreover, some of them show some largely disjunct geographic distributions on both sides of their native Palearctic region. In this study, we used a phylogenomic approach including representative populations from the oriental and occidental distribution range of B. sylvaticum micro-taxa to elucidate their evolutionary relationships and assess the systematic value of the morphological features that separate them. A combined plastome and nuclear phylogenetic tree supports an early split and high divergence of the oriental lineage, showing the close relationship of the Himalayan B. sylvaticum var. breviglume lineages to the Pacific B. miserum / B. kurilense clade, and the contrasting large homogeneity and low divergence of the occidental European, N African and SW and C Asian lineage, with several B. sylvaticum s. s., B. spryginii, and B. glaucovirens samples showing identical or similar sequences. Divergence time estimate analysis suggests that the oriental lineage diverged from the common ancestor in the early Pleistocene (2.0 Ma), followed by subsequent colonization and isolations in the Himalayas (2.0 – 1.7 Ma) and the Far East (0.36 Ma) in more recent times, while the occidental lineage split in the Mid-Late Pleistocene (0.97 Ma), followed by rapid radiation and postglacial spread in the western Paleartic during the last thousand years.