Evolutionary Relationships and Biogeography of the Ant-Epiphytic Genus Squamellaria (Rubiaceae: Psychotrieae) and Their Taxonomic Implications

Ecological research on ant/plant symbioses in Fiji, combined with molecular phylogenetics, has brought to light four new species of Squamellaria in the subtribe Hydnophytinae of the Rubiaceae tribe Psychotrieae and revealed that four other species, previously in Hydnophytum, need to be transferred to Squamellaria. The diagnoses of the new species are based on morphological and DNA traits, with further insights from microCT scanning of flowers and leaf delta C-13 ratios (associated with Crassulacean acid metabolism). Our field and phylogenetic work results in a new circumscription of the genus Squamellaria, which now contains 12 species (to which we also provide a taxonomic key), not 3 as in the last revision. A clock-dated phylogeny and a model-testing biogeographic framework were used to infer the broader geographic history of rubiaceous ant plants in the Pacific, specifically the successive expansion of Squamellaria to Vanuatu, the Solomon Islands, and Fiji. The colonization of Vanuatu may have occurred from Fiji, when these islands were still in the same insular arc, while the colonization of the Solomon islands may have occurred after the separation of this island from the Fiji/Vanuatu arc. Some of these ant-housing epiphytes must have dispersed with their specialized ants, for instance attached to floating timber. Others acquired new ant symbionts on different islands.