Trait convergence and diversification arising from a complex evolutionary history in Hawaiian species of Scaevola

Species variation in functional traits may reflect diversification relating to convergence and/or divergence depending on environmental pressures and phylogenetic history. We tested trait-environment relationships and their basis in finer-scale evolutionary processes among nine extant Hawaiian species of Scaevola L. (Goodeniaceae), a taxon with a complex history of three independent colonizations by different phylogenetic lineages, parallel ecological specialization, and homoploid hybridization events in Hawai'i. Using a wild population for each species, we evaluated traits related to plant function (morphology, leaf and wood anatomy, nutrient and carbon isotope composition). Hawaiian Scaevola species were distributed across coastal, dry forest and wet forest environments; multivariate environmental analysis using abiotic and biotic factors further showed that species from distantly related lineages inhabited similar environments. Many traits correlated with environment (based on the multivariate environmental analysis), considering both distantly related species and more closely related species. Scaevola species within shared habitats generally showed trait convergence across distantly related lineages, particularly among wet forest species. Furthermore, trait diversification through divergence was extensive among closely related Scaevola species that radiated into novel environments, especially in plant morphology and traits affecting water relations. Homoploid hybrid-origin species were "intermediate" compared to their ancestral parent species, and possessed trait combinations relevant for their current habitat. The diversity in functional traits reflected strong influences of both ecology and evolutionary history in native Hawaiian Scaevola species, and trait correspondence with environment was due to the combination of multiple processes within the taxon: trait pre-adaptation and filtering, evolutionary convergence, divergence, and hybridization.