Variation in Contact Zone Dynamics Between Two Species of Topminnows, \u3ci\u3eFundulus notatus\u3c/i\u3e and \u3ci\u3eF. olivaceus\u3c/i\u3e, Across Isolated Drainage Systems

Spatially variable selection pressure within heterogeneous environments can result in the evolution of specialist phenotypes that facilitate co-occurrence of closely related species and limit genetic exchange. If divergent selection pressures maintain reproductive isolation, hybridization is expected to correlate with the strength of underlying ecological gradients and the traits shaped by adaptive processes. We sampled ten replicate topminnow (Fundulus olivaceus and Fundulus notatus) hybrid zones in isolated drainages throughout central and southern North America. In all drainages, species were distributed in an upstream–downstream manner with contact zones localized at confluences featuring abrupt shifts from tributary to river habitat. In two drainages, the typical up and downstream positions of species were reversed. Phenotype differences between the species reflect predicted selection differences along stream gradients. Downstream populations (lower food availability and greater predator pressure) generally showed larger investment in reproduction (higher gonadal somatic index), smaller body size and lower somatic condition compared to upstream populations. Phenotypic differences between the species in the two reversed drainages were consistent with convergence of life history traits in the respective habitats. Phenotypes of individuals of hybrid origin (F1 hybrids or backcrosses) were not significantly different from the average of the two parental forms, though there were trends towards reduced fitness. The prevalence of hybridization among drainages ranged from no hybrids in two drainages to near random mating. The strongest correlates of hybridization rate among replicate hybrid zones were similarity in body shape and the homogeneity of habitat through tributary-river confluences. The two reversed orientation hybrid zones also exhibited high prevalence of hybrids suggesting that phenotypic convergence could lead to increased hybridization