The multidimensional spectrum of eco-evolutionary relationships between sharks and remoras

Remoras are a highly specialised group of fishes known to associate with a range of marine megafauna, including elasmobranchs, cetaceans and marine reptiles. Remoras appear to benefit from these interspecific interactions through consumption of host dermal parasites or reduced cost of transport. Shark-remora associations are widely documented, yet our understanding of the costs and benefits involved in these interactions is poor. Studies frequently make claims about mutualistic, commensalistic or parasitic relationships without providing the necessary quantitative information necessary to make these claims. Here I explain why this approach is problematic, and proceed to examine shark-remora interactions in a rigorous eco-evolutionary framework. These interactions cannot be properly classified without considering net evolutionary fitness and context dependence. In reality, shark-remora interactions are best defined by a multidimensional spectrum of fitness consequences, with net fitness outcomes shifting between mutualism and parasitism (and vice versa) through space and time.