First evidence of introgressive hybridization of apple snails ( spp.) in their native range.

Genetic variation facilitates both natural range expansions and anthropogenic invasions. Contrary to expectations, hybridization does not always impact negatively on biodiversity. Increasing evidence indicates advantageous roles for introgressive hybridization in maintaining standing genetic variation. Hypothesizing that hybridization may contribute to the evolutionary and invasive success of a diverse group of freshwater snails (Ampullariidae, commonly known as apple snails), we estimated the frequency of hybridization between two globally invasive species of , (Lamarck, 1822) and Perry, 1810, in their native range. While previous work in Asia has uncovered the occurrence of extensive hybridization, we provide the first phylogenetic evidence of a high degree of hybridization (30%) between these species in Uruguay and Brazil. Hybrids carried both heterozygous and homozygous combinations of elongation factor 1-α (EF1α) nuclear alleles in both mating directions, indicating that hybridization has occurred over multiple generations and likely preceded introductions outside the native range. Among the five sites in Brazil previously documented as containing only , one far northern population (Careiro Castanho), which is thousands of kilometres from the northern range of , unexpectedly contained hybrids. This may be the result of human-facilitated introductions. Together with recent work from Asia, our investigations in the native range of apple snails support a reframing of historical perspectives of hybridization as a driver of extinction and diversity loss towards a modern paradigm where hybridization may promote diversification and contribute to the survival of evolutionary lineages such as molluscs.