Manipulated sex ratios alter group structure and cooperation in the brown-headed nuthatch

A biased adult sex ratio (ASR) can influence cooperative breeding behavior if the bias limits mating opportunities for the more abundant sex. We tested predictions associated with the ASR-cooperation hypothesis in the brown-headed nuthatch (Sitta pusilla). We manipulated ASR by cross-fostering known-sex nestlings within 2 large (≥100 ha) experimental plots for 5 years using a crossover design where each plot received an opposing male- or female-biased treatment for 2 consecutive years. A year with no manipulations followed before the bias was reversed on each plot for 2 additional years. Variation in ASR (adult males/total adults) was pronounced compared to background proportions (0.55) and ranged from a female bias in female-biased plots (0.47) to a strong male bias in male-biased plots (0.71). Sex ratios during the postbreeding period ranged more broadly (0.33 in female-biased plots vs. 0.74 in male-biased plots). Territory densities did not change significantly and allowed 6 predictions to be assessed. Consistent with predictions, the prevalence of cooperative breeding groups doubled under male-biased treatments and large cooperative groups appeared (≥2 male helpers vs. the single male helper most common prior to the experiment). These changes occurred despite increased dispersal of cross-fostered males in male-biased plots. Most juvenile females dispersed, but, consistent with predictions, the prevalence of female helpers increased under female-biased treatments. Manipulations did not alter the sex of nestlings produced nor extend the time that males served as helpers. Taken collectively, results support the ASR-cooperation hypothesis and the role that mate limitations play in cooperative breeding behavior.