We unfold the bifurcation involving the loss of evolutionary stability of an equilibrium of the canonical equation of Adaptive Dynamics (AD). The equation deterministically describes the expected long-term evolution of inheritable traits — phenotypes or strategies — of coevolving populations, in the limit of rare and small mutations. In the vicinity of a stable equilibrium of the AD canonical equation, a mutant type can invade and coexist with the present — resident — types, whereas the fittest always win far from equilibrium. After coexistence, residents and mutants effectively diversify, according to the enlarged canonical equation, only if natural selection favors outer rather than intermediate traits — the equilibrium being evolutionarily unstable, rather than stable. Though the conditions for evolutionary branching — the joint effect of resident-mutant coexistence and evolutionary instability — have been known for long, the unfolding of the bifurcation has remained a missing tile of AD, the reason being related to the nonsmoothness of the mutant invasion fitness after branching. In this paper, we develop a methodology that allows the approximation of the invasion fitness after branching in terms of the expansion of the (smooth) fitness before branching. We then derive a canonical model for the branching bifurcation and perform its unfolding around the loss of evolutionary stability. We cast our analysis in the simplest (but classical) setting of asexual, unstructured populations living in an isolated, homogeneous, and constant abiotic environment; individual traits are one-dimensional; intra- as well as inter-specific ecological interactions are described in the vicinity of a stationary regime.
Adaptive dynamics of unstable cancer populations: The canonical equation
