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A resident trait $x$ occupies fraction $1-m/c(x)$ when its colonization rate exceeds fire extinction. With net competitive replacement given by ability difference $B(y)-B(x)$, a rare mutant has the displayed per-capita growth. The selection gradient is $g(x)=m{c}^{\prime }(x)/c(x)+[1-m/c(x)]B^{\prime }(x)$. A singular trait with negative mutant-fitness curvature is a strict local evolutionarily stable strategy. Global strict uninvadability requires that $mc(y)+[c(x)-m]B(y)$ be smaller than its resident value for every other trait. Monotonicity alone does not specify the shape or existence of an optimum.

Let $a,b,e$ be fractions of patches occupied by the two competitors and empty, with $a+b+e=1$. Assume well-mixed propagule dispersal, equal independent fire extinction rate $m>0$, colonization constants $c_A>c_B>0$, and net replacement advantage $q=k_B-k_A>0$ for the poorer colonizer. The Levins metapopulation model with replacement isReplacement fluxes $k_{B}ab$ and $k_{A}ab$ cancel in the total occupancy equation. At every face the outward flux is zero or points into the patch probability simplex, so these equations preserve valid fractions. The constants include both propagule arrival and establishment probability; the model does not require unequal fire susceptibility.

Put $\Delta c=c_A-c_B$. Solving the two occupied-patch balances and their sum constraint givesThus strict coexistence is equivalent toToo little disturbance lets the better competitor exclude the colonizer; too much disturbance favors colonization or destroys occupancy. A positive equilibrium automatically has $m<c_A$. If the poorer colonizer cannot survive alone, coexistence can nevertheless be possible because it repeatedly replaces established patches of the better colonizer.

These bounds also have an invasion interpretation. The $A$-only equilibrium is $a=1-m/c_A$ when $c_A>m$; the growth rate of rare $B$ there is $q-m(q+\Delta c)/c_A$, positive at the upper-bound condition above. At a feasible $B$-only equilibrium the rare-$A$ growth is $-q+m(q+\Delta c)/c_B$, giving the lower bound. At coexistence the occupied-patch Jacobian matrix has trace $-c_A{a}_{\ast }-c_B{b}_{\ast }<0$ and determinant $a_{\ast }{b}_{\ast }q(q+\Delta c)>0$, proving local asymptotic stability. The Bendixson-Dulac criterion with multiplier $1/(ab)$ gives divergence $-c_A/b-c_B/a<0$ in the interior, excluding periodic competition cycles.

To specify selection, a monotone tradeoff alone is insufficient: one must say how two different trait values determine replacement. A natural model uses a trait $x$ with increasing competitive ability $B(x)$ and decreasing colonization $c(x)$, and net replacement $B(y)-B(x)$ by a mutant $y$ against a resident $x$. The resident metapopulation occupies $1-m/c(x)$, requiring $c(x)>m$. The rare-mutant per-capita growth, or invasion fitness, isIt is zero at $y=x$. Differentiating with respect to the mutant trait gives the selective pressureHigher colonization is valuable in empty habitat; higher competitive ability is valuable where the resident already occupies patches. This is colonization-competition tradeoff invasion fitness.

At an interior candidate for a local evolutionarily stable strategy, $g(x_{\ast })=0$. A strict, nondegenerate local evolutionarily stable strategy follows ifsince nearby distinct mutants then have negative invasion fitness. For a strict global evolutionarily stable strategy, every feasible distinct mutant must satisfyThis is the supporting-line condition on the tradeoff curve. At a trait boundary replace the vanishing-gradient condition by the appropriate one-sided inequality. Equality against another mutant, or zero local curvature, needs the second condition of evolutionary stability or higher-order terms; the strict tests above are sufficient and are the usual nondegenerate conditions, not a claim that every tie is unstable.

For evolutionary approach to a singular point, the selection gradient must also point toward it, locally $g^{\prime }(x_{\ast })<0$. This is logically distinct from testing mutant fitness. In this particular additive-ability model with $c^{\prime }<0<B^{\prime }$, negative mutant curvature implies negative $g^{\prime }$ becauseIf replacement instead has a general kernel $\kappa (y,x)$, replace $B(y)-B(x)$ in $f$ by that kernel and differentiate it; monotonicity does not justify importing the displayed additive formula without that modeling assumption. The tradeoff shape, disturbance rate and replacement law together determine selection and local or global uninvadability.

For a differentiable rare-mutant invasion fitness $f(y,x)$ against resident trait $x$, the selection gradient gives the first-order growth advantage of a nearby mutant. A zero gradient is a candidate interior evolutionary singularity. Strict local uninvadability follows from negative mutant-fitness curvature; evolutionary approach additionally requires the gradient to point toward the candidate. Boundary traits require one-sided tests. A gradient alone does not prove global evolutionary stability.