Sympatric speciation under incompatibility selection.

The existing theory of sympatric speciation assumes that a local population splits into two species under one-dimensional disruptive selection, which favors both of the opposite extreme values of a quantitative trait. Here we model sympatric speciation under selection that favors high values of either of the two independently inherited traits, each required to efficiently consume one of the two available resources, but acts, because of a tradeoff, against those possessing high values of both traits. Such two-dimensional incompatibility selection is similar to that involved in allopatric speciation. Using a hypergeometric phenotypic model, we show that incompatibility selection readily leads to sympatric speciation. In contrast to disruptive selection, two distinct modes of sympatric speciation exist under incompatibility selection: under strong tradeoffs both of the new species are specialists, each consuming its own resource, but under moderate tradeoffs speciation may be asymmetric and involve the origin of a specialist and a generalist species. Also, incompatibility selection may lead to irreversible specialization: under strong tradeoffs, the population speciates if it consists mostly of unspecialized individuals, but remains undivided if most of the individuals are specialized to consume one of the resources. Incompatibility selection appears to be more realistic than disruptive selection, implying that incompatibility between individually adaptive alleles or trait states drives both allopatric and sympatric speciation.