Evolutionary responses to conditionality in species interactions across environmental gradients

The outcomes of many species interactions are conditional on the environments in which they occur. A common pattern is that outcomes grade from being more positive under stressful conditions to more antagonistic or neutral under benign conditions. The evolutionary implications of conditionality in interactions have received much less attention than the documentation of conditionality itself, with a few notable exceptions. Here, we predict patterns of adaptation and co-adaptation between partners along abiotic gradients, positing that when interactions become more positive in stressful environments, fitness outcomes for mutations affecting interactions align across partners and selection should favor greater mutualistic adap-tation and co-adaptation between interacting species. As a corollary, in benign environments, if interactions are strongly antagonistic, we predict antagonistic co-adaptation resulting in Red Queen or arms-race dynamics, or reduction of antagonism through character displacement and niche partitioning. We predict no adaptation if interactions are more neutral. We call this the CoCoA hypothesis: ( Co )-adaptation and Co nditionality across A biotic gradients. We describe experimental designs and statistical models that allow testing predictions of CoCoA, with a focus on positive interactions. While only one study has included all the elements to test CoCoA, we briefly review the literature and summarize study findings relevant to CoCoA and highlight opportunities to test CoCoA further.