Don't ask “when is it coevolution?”—ask “how?”

Coevolution has come to be widely understood as specific, simultaneous, reciprocal adaptation by pairs of interacting species. This strict-sense definition arose from a desire for conceptual clarity, but it has never reflected the much wider diversity of ways in which interacting species may shape each other's evolution. As a result, much of the literature on the evolutionary consequences of species interactions pays homage to the strict-sense definition while addressing some other form of coevolution. This tension suggests we should reframe the key question in coevolution research, from "when is it coevolution?" to, rather, "how is it coevolution?". The result is not so much a definition of coevolution as a mission statement: We can describe how species coevolve by documenting the ways that each species shaped the other's genetic diversity over a shared history of interaction. Making this change shifts our focus from identifying case studies for a single, narrowly defined process to describing the many ways-specific and diffuse, simultaneous and stepwise, and adaptive and nonadaptive-in which species evolve together.