Interacting selection diversifies warning signals in a polytypic frog: an examination with the strawberry poison frog

Aposematic signals represent one of the most accessible traits to evaluate the interaction of natural and sexual selection on signal evolution. Here we investigate the contributions of these two selective forces on the aposematic signal evolution of the highly polytypic strawberry poison frog, Oophaga pumilio, of Bocas del Toro, Panama. Previous research has shown that the brightness of O. pumilio warning coloration can inform predators of the toxicity levels associated with different populations of the archipelago. Other studies suggest that sexual selection may be influencing warning signal brightness within populations via female mate choice (Isla Solarte, Isla Bastimentos, and Aquacate Peninsula populations) and male-male competition (Isla Solarte). Here we present two non-exclusive scenarios for how natural and sexual selection interact to drive phenotypic variation across this archipelago: (1) predators impose a selective regime whereby populations above a toxicity-brightness threshold are at liberty to diversify via sexual selection and below which populations are constrained to maintain a stricter resemblance to a more cryptic population mean, and (2) synergistic/additive effects of inter- and intrasexual selection drive the evolution of brighter males within populations above this toxicity threshold. We investigate whether aposematic patterns of divergence across the archipelago relative to the common mainland phenotype meet these predictions using existing data on O. pumilio morph toxicity measures and overall conspicuousness estimates to an avian predator. Using standardized z-scores to evaluate the range of trait values, we find that indeed the population representative of the common mainland phenotype (Almirante) represents an intermediate level of both toxicity and conspicuousness, and that derived Bocas del Toro populations vary in each of those components in directions predicted by the proposed scenarios. Furthermore, we find greater divergence towards conspicuousness than crypsis, a pattern suggestive of sexual and natural selection acting synergistically in morphs with high toxicity.