Gene expression bias is associated with directional differentially methylated genes (DMGs) in S. invicta. DMGs that exhibit elevated methylation in haploid males are more highly expressed in haploid males, whereas DMGs that exhibit elevated methylation in diploid (a) males, (b) queens or (c) workers are more highly expressed in diploid males, queens or workers, respectively. By contrast, there is no significant difference between the ratio of expression for DMGs that exhibit elevated methylation in (d) a pairwise comparison of queens and workers. Expression ratio data were standardized (mean zero, unit variance) following log2-transformation; p-values denote the results of Mann–Whitney U-tests. (Online version in colour.)