Phylogenomics of Brosimum (Moraceae) and allied genera, including a revised subgeneric system

We present a phylogenomic study of Brosimum (Moraceae) and the allied genera Trymatococcus and Helianthostylis, with near-complete taxon sampling. Distributed from Mexico and the Greater Antilles to the Amazon, this clade contains the underutilized crop ramon (bread nut) (Brosimum alicastrum) as well as other species valued for timber or medicinal uses. Target enrichment for 333 genes produced a well-resolved phylogenetic tree and showed that Trymatococcus and Helianthostylis are nested within Brosimum. We present a revised subgeneric classification of Brosimum (19 spp.) based on phylogenetic and morphological considerations, including the reduction of Trymatococcus and Helianthostylis to subgenera. The monophyletic subgenera can be diagnosed based on stipule, pistillode, and cotyledon synapomorphies. Divergence date estimates suggest a Miocene origin for Brosimum, and ancestral area reconstruction indicated that all four subgenera originated and initially diversified in Amazonia before dispersing into other parts of South and Central America.