Is sexual brutality maladaptive under high population density?

Coercive copulation is frequent in promiscuous mating systems, generating both benefits (e.g. higher copulation rate) and costs (e.g. injuries and fewer foraging opportunities). The negative consequences of sexual harassment are expected to increase with increasing population density with male-biased operational sex ratio (OSR). This study offers an example in which the frequency and severity of the injuries inflicted on female tortoises during forced copulations increased with population density and biased OSR. Male Hermann’s tortoises (Testudo hermanni) harass females during courtship, ramming and biting them until they eventually force mating. They also use their horny and mobile tail to stimulate females, sometimes damaging their cloaca. We compared the frequency and severity of cloacal injuries between two populations in the Republic of Macedonia. Under high density and balanced OSR at one site (Konjsko), 25% of females exhibited cloacal injuries. Under very high density and extremely biased OSR at another site (Golem Grad Island), 75% of females were wounded, often severely. Furthermore, exclusively on Golem Grad, many small and immature females were courted and wounded. This behavioural shift toward juveniles generated costs without benefits. Perhaps brutal reproductive behaviours in male Hermann’s tortoises evolved under low or moderate population densities? Under very high densities, coercive mating seems maladaptive, negatively affecting females and possibly further skewing OSR.