Male-killingWolbachiain a flour beetle

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Abstract
The bacteria in the genus Wolbachia are cytoplasmically inherited symbionts of arthropods. Infection often causes profound changes in host reproduction, enhancing bacterial transmission and spread in a population. The reproductive alterations known to result from Wolbachia infection include cytoplasmic incompatibility (CI), parthenogenesis, feminization of genetic males, fecundity enhancement, male killing and, perhaps, lethality. Here, we report male killing in a third insect, the black flour beetle Tribolium madens, based on highly female–biased sex ratios of progeny from females infected with Wolbachia. The bias is cytoplasmic in nature as shown by repeated backcrossing of infected females with males of a naturally uninfected strain. Infection also lowers the egg hatch rates significantly to approximately half of those observed for uninfected females. Treatment of the host with antibiotics eliminated infection, reverted the sex ratio to unbiased levels and increased the percentage hatch. Typically Wolbachia infection is transmitted from mother to progeny, regardless of the sex of the progeny; however, infected T. madens males are never found. Virgin females are sterile, suggesting that the sex–ratio distortion in T. madens results from embryonic male killing rather than parthenogenesis. Based on DNA sequence data, the male–killing strain of Wolbachia in T. madens was indistinguishable from the CI–inducing Wolbachia in Tribolium confusum, a closely related beetle. Our findings suggest that host–symbiont interaction effects may play an important role in the induction of Wolbachia reproductive phenotypes.