In the two-class globally coupled underdominant metapopulation, the rectangle , is forward invariant under the displayed bound. On its inner edges, selection points inward with magnitude while outward migration is at most or ; the outer edges also point inward. Thus prescribed initial state cannot approach global allele fixation when . Maximizing gives at , making the bound optimal within this particular symmetric worst-case rectangle argument, not a universal sharp migration threshold.
Use a globally coupled underdominant metapopulation with equal habitat sizes:
Symmetry preserves a common frequency in the initially habitats and in the initially habitats. With , the equations reduce exactly to
The maximum positive selection rate is at . Comparing it with the worst outward migration flux gives a sufficient barrier condition . A slightly stronger optimized condition follows from maximizing the per-capita restoring term over : its maximum is at .
Indeed, on , with , . On , with , . The other two edges point inward as well. The invariant-rectangle criterion for underdominant coexistence therefore gives the explicit robust conditions
Starting from the specified pure habitats, the populations then remain in , . Their global frequency lies between and , so neither allele fixes. This is a sufficient condition, not a claim that the bound is sharp. If or , the population is already fixed; if , any mixture of the two pure habitat types persists.
The exactly balanced case illustrates why robustness and the prescribed trajectory differ. When , symmetry forces and
For its heterogeneous limiting state is
Here , so the full Jacobian matrix has diagonal entries and off-diagonal entries . Its two full-system eigenvalues are and : the symmetric underdominant two-cluster equilibrium is robustly attracting only for . For larger migration the exactly symmetric initial condition can still avoid fixation, eventually approaching the spatially uniform frequency when , but the symmetry-broken mean-frequency mode is unstable. A finite population with genetic drift need not maintain deterministic coexistence indefinitely.