The evolutionary terrestrialization of planarian flatworms (Platyhelminthes, Tricladida, Geoplanidae): a review and research programme

The terrestrialization of animal life from aquatic ancestors is a key transition during the history of life. Planarian flatworms form an ideal group of model organisms to study this colonization of the land because they have freshwater, marine, and terrestrial representatives. The widespread occurrence of terrestrial flatworms is a testament to their remarkable success occupying a new niche on land. This lineage of terrestrial worms provides a unique glimpse of an evolutionary pathway by which a group of early divergent aquatic, invertebrate metazoans has moved onto land. Land flatworms are among the first groups of animals to have evolved terrestrial adaptations and to have extensively radiated. Study of this terrestrialization process and the anatomical key innovations facilitating their colonization of the land will contribute greatly to our understanding of this important step in Metazoan history. The context and scientific background are reviewed regarding the evolutionary terrestrialization of land flatworms. Furthermore, a framework of a research programme is sketched, which has as its main objective to test hypotheses on the evolution of land planarians, specifically whether particular anatomical and physiological key innovations have contributed to their evolutionary successful terrestrial colonization and radiation. In this context special attention is paid to the respiration in aquatic and terrestrial planarians. The research programme depends on a comprehensive phylogenetic analysis of all major taxa of the land flatworms on the basis of both molecular and anatomical data. The data sets should be analyzed phylogenetically with a suite of phylogenetic inference methods. Building on such robust reconstructions, it will be possible to study associations between key innovations and the evolutionary terrestrialization process.