Sexual dimorphism of body size and shell shape in European tortoises

Adult body size and shape were examined in almost 1400 individuals of the tortoises Testudo graeca , T. hermanni and T. marginata from Greece. The size at maturity was greater in females than in males in all three species. Maximum and mean adult sizes were also greater in females than in males in T. graeca and T. hermanni . Males grew to a larger size than females in T. marginata , and mean adult size was similar in the sexes in this species. Sexual dimorphism of shape (adjusted for size covariate) was shown in most of the characters examined, and the degree of this dimorphism differed significantly among the three species. Differences were related to their contrasting courtship behaviours: horizontal head movements and severe biting in T. marginata , vertical head bobs and carapace butting in T. graeca , and mounting and tail thrusting in T. hermanni . There was no difference in the frequency of observations of courtship or fighting among the three species, but courtship was about 10 times more common than combat in males. All species showed greatest courtship activity in autumn; copulation was rarely observed in T. hermanni (only 0.36% of courting males) and not seen in the other species in the field. Observations made throughout the activity season indicated that feeding was equally common in males and females in all three species. Differences in shape were more likely to be the result of sexual selection than of natural selection for fecundity. Detailed predictions are made for sexual dimorphism of other characters in these species.