Haplodiploidy and the Evolution of Facultative Sex Ratios in a Primitively Eusocial Bee

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Abstract
In eusocial Hymenoptera, the haplodiploid system of sex determination creates relatedness asymmetries such that workers are more closely related on average to their sisters than to their brothers. For such societies, kin-selection theory and sex-ratio theory predict that workers maximize their inclusive fitness by biasing the investment sex ratio toward females. To test the prediction of sex-ratio biasing, relatedness asymmetries were experimentally manipulated in colonies of the primitively eusocial bee Augochlorella striata (Halictidae: Hymenoptera) by removing or not removing foundress queens. Queenright colonies (relatedness asymmetry present) produced a more female-biased sex ratio than did queenless colonies (relatedness asymmetry absent). Worker reproduction and unmated replacement queens can be discounted as alternative explanations. Workers therefore facultatively adjusted their colony's sex ratio and, in the presence of a relatedness asymmetry, biased the investment sex ratio toward their more closely related sisters and away from their more distantly related brothers.

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