Haldane rules: Costs of outbreeding at production of daughters in sand lizards

Haldane's rule is one of the most widely applicable paradigms in evolutionary biology, stating that in species crossings, the heterogametic sex will suffer more severely in terms of sterility and inviability. We address this in a within-species outbreeding situation by assessing the risk of producing inviable offspring depending on the sex ratio of the clutch produced in between-population crossings in the laboratory. In crossings between male and female sand lizards (Lacerta agilis) from two different sampling regions, one in Sweden, one in central Europe, risk of gametic incompatibility is unaffected by outbreeding, but offspring from between-population crossings show 300% higher malformation frequency and 10% lower hatching success. The risk of having in viable offspring increases with the production of daughters, i.e. the hemizygous sex in this species (ZW). Such sex-specific genetic costs of offspring production need to be incorporated into life history ecology, e.g. sex allocation theory.