Phylogeny of Coussareeae (Rubioideae, Rubiaceae)

Coussareeae are a Neotropical clade of morphologically heterogeneous plants in the subfamily Rubioideae of the coffee family (Rubiaceae). The tribe encompasses about 330 species assigned to ten genera: Bradea, Coccocypselum, Coussarea, Cruckshanksia, Declieuxia, Faramea, Heterophyllaea, Hindsia, Oreopolus, and Standleya. Historically, the genera of Coussareeae have rarely been considered closely related, and the widely defined Coussareeae were delimited based on molecular systematics without proposed morphological synapomorphies. In order to assess the tribe’s monophyly, as well as the generic limits, infrageneric relationships, and suprageneric relationships, multiple specimens per genus were sampled whenever possible and analyzed using multiple molecular loci with the Bayesian and maximum likelihood methods. The results of the phylogenetic analyses (all genera represented by multiple terminals are monophyletic, all genera are resolved with respect to each other, and three major suprageneric clades are resolved), coupled with herbarium and literature studies, were used to identify potential synapomorphic features. Non-molecular diagnostic features remain elusive for Coussareeae as a whole, but we have identified multiple diagnostic features and potential synapomorphies for each of the three major suprageneric clades: (1) Coussarea and Faramea (e.g., porate pollen grains with annuli bordering the pores); (2) Bradea, Coccocypselum, Declieuxia, Hindsia, and Standleya (e.g., colporate pollen grains with complex reticulate tecta); and 3) Cruckshanksia, Heterophyllaea, and Oreopolus (e.g., chartaceous, loculicidal capsules). The latter clade, distributed in different biomes of the Andes, is sister to the former two, both widely distributed in the Neotropics.