Proximate and ultimate control of sex ratios inMyrmica brevispinosacolonies

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Abstract
The literature on sex ratio evolution in ant colonies is dominated by inclusive fitness arguments. In general, genetic theory makes good predictions about sexual investment in ant populations, but understanding colony–level variance in sex investment ratios has proven more difficult. Recently, however, more studies have addressed ecological factors that influence colony–level sex investment ratios. Food availability, in particular, has been manipulated because larval nutrition influences female caste determination, thus implying that resource availability should be of critical importance for colony sex investment ratios. However, results from food supplementation experiments are equivocal, and it is clear that ant colony response to food supplementation is dependent on the ecological background of the population. We presented field colonies of the ant Myrmica brevispinosa with two food types (proteins and carbohydrates), and assessed their relative impact on colony–level sex investment ratios. We show that colonies receiving carbohydrate enhancement invested in more female sexuals and produced more female–biased sex allocation ratios than protein–fed or control colonies. Thus, our study is the first, to our knowledge, to demonstrate that sex ratios in social insect colonies might be sensitive to resource quality. Male investment was not influenced by food treatment, but was positively correlated with colony size. Therefore, the shift in sex ratios in our study must have been mediated through nutritional influences on female caste determination rather than male brood elimination. We also used our data to evaluate evidence for sex ratio compensation by queenright colonies in response to male production by workers from orphaned colonies.