Evolutionary mode, tempo, and phylogenetic association of continuous morphological traits in the aquatic moss genus Amblystegium

Evolutionary significance of morphological characters that have traditionally been used for species delineation in the aquatic moss genus Amblystegium was tested by partitioning the environmentally and genetically induced morphological variation and focusing on morphological evolution using comparative methods. Cultivation experiments under controlled condition showed that most of the morphological variation in nature resulted from plasticity. Information regarding genetically fixed morphological variation and genetic similarity derived from polymorphic inter-simple sequence repeat markers was combined into an explicit model of morphological evolution. Maximum likelihood estimates of the model parameters indicated that evolution of most characters tended to accelerate in the most recent taxa and was often independent from the phylogeny. Constraining the different characters to be independent from each other most often produced a less likely result than when the characters were free to evolve in a correlated fashion. Thus, the morphological characters that have traditionally been used to circumscribe different Amblystegium species lack the independence, diagnostic value for specific lineages, and stability that would be required for distinguishing different species.