Making the grade: Physiological adaptations to terrestrial environments in decapod crabs

All extant macroscopic terrestrial diversity has evolved from the ancestors of a small group of successful terrestrial colonizers, but in a few lineages this transition has independently occurred multiple times in spite of the significant functional challenges it presents. Decapod crabs have transitioned from marine to terrestrial environments at least ten times, occupy diverse habitats, and display varying degrees of terrestriality. Previous attempts to categorize land crab diversity relied on single traits, did not explicitly distinguish between brachyuran and anomuran lineages, and did not separate lineages that colonized land via freshwater or marine environments. As a result, critical phylogenetic and ecological constraints were missing from these earlier classifications. In this paper, I reclassify terrestriality in the land crabs by designating four transition pathways that reflect deep phylogenetic relationships between the two decapod crab infraorders and the route-specific nature of this transition. I then describe the adaptive traits that evolved in response to six primary terrestrial selective challenges. I conclude by proposing six grades of terrestriality in this system that describe observable trait-by-environment associations, and propose studies that can test the hypothetical sequence of trait evolution and the nature of convergence in the land crabs using phylogenomic and transcriptomic tools.