Evolutionary history, ecological divergence, and introgression in the Oncocyclus irises species complex in the Southern Levant

Speciation is a dynamic process shaped by the interaction of gene flow, geographic isolation, and ecological divergence. The Oncocyclus irises of the Southern Levant represent a young radiation of narrowly endemic species considered to be in the course of speciation. In this study, we used RAD-sequencing and single nucleotide polymorphism analysis across nine described species to investigate patterns of genomic divergence, gene flow, and local adaptation. Phylogenomic analyses revealed a mix of well-supported clades for some species, previously defined by their morphology and distribution, and non-monophyletic lineages, with several species exhibiting shallow divergence and shared genetic ancestry. We found evidence for gene flow and historical introgression between Iris petrana, I. atrofusca and I. mariae, while other cases of non-monophyly appear driven by incomplete lineage sorting. Both geographic distance (IBD) and environmental factors (IBE), mainly altitude, temperature, and aridity, were significantly associated with genetic structure, suggesting that local adaptation contributed to divergence following range expansion. Based on our findings, we propose that the divergence of the Oncocyclus iris species in the Southern Levant supports a stepping-stone dispersion model, in which north-to-south dispersal was followed by local adaptation, and introgression in secondary contact zones. Overall, these findings highlight the complexity of speciation and the need for integrative approaches to study the interplay between historical divergence, contemporary gene flow, and environmental differentiation in shaping genomic patterns.