Experimental evidence for habitat dependent selection in a Bombina hybrid zone

Hybridizing taxa remain distinct for two main reasons. Natural selection acts against hybrids either because of their incompatible genome, or because of differential adaptation of the pure types across an environmental gradient. Here, we provide experimental evidence that the location of the Bombina (Anura: Discoglossidae) hybrid zone in Croatia is, at least in part, determined by differential adaptation. B. bombina typically breeds in permanent water in the lowland, whereas B. variegata reproduces in puddles at higher elevations. In a reciprocal translocation, pure bombina and variegata tadpoles were introduced in equal proportions into lowland pond enclosures and upland puddles. After three weeks, variegata exceeded bombina in survival and growth in both habitats. The effect was most pronounced in puddles, where the few surviving bombina tadpoles had hardly grown at all. In comparison to variegata, the smaller hatchlings of bombina grew relatively faster in ponds, but remained smaller in absolute terms. Nevertheless, B. bombina appears better adapted to ponds than to puddles. The mechanisms by which variegata is excluded from ponds remain to be demonstrated. These data show that habitat dependent selection prevents the invasion of bombina tadpole traits into the variegata gene pool. Given the strong linkage disequilibria in hybrid populations, differential selection on tadpoles may be sufficient to maintain the integrity of the two gene pools.