Adult survival and recruitment and the explanation of an uneven sex ratio in a tortoise population

Survival, recruitment, and dynamics of adult Testudo hermanni at Alyki (northern Greece) were studied from 1980 to 1988. Recruitment of adult males was greater than that of females owing to their shorter time to maturity (9 vs. 11 years); recruitment of subadults (6 years old) was equal in males and females. Mean annual survival was slightly greater in males (0.914) than females (0.877), equivalent to mean adult longevity values of 11.6 and 8.1 years, respectively. Excluding tortoises that die before maturity, male and female T. hermanni are mature for about 56 and 42% of their life, respectively. Generation time was roughly twice the age at maturity, and three times the age at which secondary sexual characters develop, a pattern which may apply to other tortoises. The combination of adult survival and recruitment should lead to a stable sex ratio (males/females) of 2.1. The adult sex ratio was higher than this, but decreasing, from4.1 in 1982to3.0 in 1986, with an increase in the female population; the number of males was stable. The observed sex ratio showed a similar decline and a further fall to 2.4 in 1988. The cause of the lower survival rate of females compared with males is proposed to be damage during courtship attempts. The courtship behaviour and related anatomy of T. hermanni are compared with those of T. graeca, a species with even population sex ratios. The level of male-induced female mortality would depend on population density; the increasing number of females during the study follows the approximate halving of population density in 1980.