Dobzhansky's rule and reinforcement of pre-zygotic reproductive isolation in zones of secondary contact

It is well known that closely related sympatric species are usually more different in characters involved in species recognition (e.g. in visual and acoustic signals) than allopatric species of the same evolutionary age. In this article I call this phenomenon Dobzhansky's rule in accordance with the name of the scientist who first discovered it. There are two alternative explanations for this pattern. Under hypothesis of reinforcement by Dobzhansky, these species-specific differences evolve in situ, exactly in zone of overlap between two populations. Under hypothesis of differential fusion by Templeton, the differences originate in geographically separated regions, and only those populations that have evolved such differences can persist in secondary sympatry. These evolutionary scenarios are significantly different. The scenario by Dobzhansky is an essentially sympatric model, in which natural selection reinforces pre-zygotic isolation between divergent populations by selecting against unfit hybrids. The scenario by Templeton is based on classic allopatric speciation model that consider the formation of reproductive isolation to be a by-product of divergent evolution. In this work we show that the sympatric distribution of sister taxa of Agrodiaetus butterflies strongly correlates with differences in male wing colour. We also use a new quantitative phylogenetic test to distinguish between the models by Dobzhansky and by Templeton and to demonstrate that the pattern observed is, most likely, the result of reinforcement.