Mutualism between antagonists: its ecological and evolutionary implications.

Mutualism or antagonism between species is often investigated within the framework of monotonic interactions of either mutualism or antagonism, but studies on transition from mutualism to antagonism (within the context of nonmonotonic interactions) have been largely ignored. In this paper, through a brief review and synthesis, we highlighted the role of mutualism between antagonists in regulating the ecological and evolutionary processes, as well as maintaining the stability and complexity of ecosystems. Mutualism between antagonistic species represents the density-dependent transition between mutualism and antagonism, which is beneficial to species coexistence and stability of complex ecosystems; thus, it should be favored by natural selection. Species may face selection of conflicting pressure on functional traits in co-balancing mutualism and antagonism, which may result in evolution of the dual character of species with moderate mutualistic or antagonistic traits. Coevolution and co-balance of these traits are driving forces in shaping mutualism-antagonism systems. Rewards for mutualists, punishment for exploiters, and competition of meta-communities are essential in stabilizing mutualism between antagonists. We appeal for more studies on mutualism between antagonists and its ecological and evolutionary implications by expanding the conventional ecological studies from monotonic to nonmonotonic regimes.