Sea lettuce systematics: lumping or splitting?

Phylogenetic relationships within sea lettuce species belonging to the genus Ulva is a daunting challenge given the scarcity of diagnostic morphological features and the pervasive phenotypic plasticity. With more than 100 species described on a morphological basis, an accurate evaluation of its diversity is still missing. Here we analysed 277 chloroplast-encoded gene sequences (43 from this study), representing 35 nominal species of Ulva from the Pacific, Indian Ocean, and Atlantic (with a particular emphasis on the Brazilian coast) in an attempt to solve the complex phylogenetic relationships within this widespread genus. Maximum likelihood, Bayesian analyses and species delimitation tests support the existence of 22 evolutionary significant units (ESUs), lumping the currently recognized number of species. All individuals sampled throughout an extensive area of the Brazilian coast were included within two distinct ESUs. Most of the clades retrieved in the phylogenetic analyses do not correspond to a single nominal species. Geographic range evolution indicated that the ancestor of Ulva had a distribution restricted to the temperate North Pacific. The dating analysis estimated its origin during the Upper Cretaceous at 75.8 million years (myr) ago but most of the cladogenetic events within the genus occurred in the last ten myr. Pervasive human-mediated gene flow through ballast water and widespread morphologic plasticity are the most likely explanations for the difficulty in establishing a reliable phylogenetic framework for this conspicuous, widespred and many times abundant green algae morphotype.