Determinants of reproductive success and offspring sex in a turtle with environmental sex determination

Abstract

Despite the importance of maternal effects in evolution, and knowledge of links among nest site choice, timing of nesting, offspring sex, and reproductive success in animals with environmental sex determination, these attributes have not been rigorously studied in a combined and natural context. To address this need we studied the relationships between three maternal traits (nest site choice, lay date, and nest depth) and two fitness-related attributes of offspring (hatchling sex and embryonic survival) in the riverine turtle Carettochelys insculpta, a species with temperature-dependent sex determination, for four years. Predation and flooding were the major sources of embryonic mortality in 191 nests. Embryonic survival was influenced by both lay date and nest site choice: in one year when nesting began later than average, nests laid later and at lower elevations were destroyed by early wet season river rises. In other years early nesting precluded flood mortality. However, turtles did not nest at the highest available elevations, and a field experiment confirmed that turtles were constrained to nest at lower elevations where they could construct a nest chamber. The principal determinant of hatchling sex in 140 nests was lay date, which in turn was apparently related to the magnitude of the previous wet season(s). Clutches laid earlier in the season (a female's first clutch) produced mainly males, while later clutches (her second clutch) yielded mostly females, due to seasonal increases in air temperatures. Accordingly, later nesting produced female-biased hatchling sex ratios in 1996, while earlier nesting resulted in sex ratios near unity in the other years. However, all-female nests were more likely to be flooded than mixed-sex or all-male nests in years when nesting was late. In conclusion, we found evidence that the position of two maternal trait distributions (elevation of the nest site and lay date), associated with the reproductive strategy of C. insculpta, reflect a combination of natural selection, physical constraints, and phenotypic plasticity.