Measurement and modelling of primary sex ratios for species with temperature-dependent sex determination.

For many oviparous animals, incubation temperature influences sex through temperature-dependent sex determination (TSD). Although climate change may skew sex ratios in species with TSD, few available methods predict sex under natural conditions, fewer still are based on mechanistic hypotheses of development, and field tests of existing methods are rare. We propose a new approach that calculates the probability of masculinization (PM) in natural nests. This approach subsumes the mechanistic hypotheses describing the outcome of TSD, by integrating embryonic development with the temperature-dependent reaction norm for sex determination. Further, we modify a commonly used method of sex ratio estimation, the constant temperature equivalent (CTE), to provide quantitative estimates of sex ratios. We test our new approaches using snapping turtles (Chelydra serpentina). We experimentally manipulated nests in the field, and found that the PM method is better supported than the modified CTE, explaining 69% of the variation in sex ratios across 27 semi-natural nests. Next, we used the PM method to predict variation in sex ratios across 14 natural nests over 2 years, explaining 67% of the variation. We suggest that the PM approach is effective and broadly applicable to species with TSD, particularly for forecasting how sex ratios may respond to climate change. Interestingly, we also found that the modified CTE explained up to 64% of variation in sex ratios in a Type II TSD species, suggesting that our modifications will be useful for future research. Finally, our data suggest that the Algonquin Park population of snapping turtles possesses resilience to biased sex ratios under climate change.

A new method of estimating thermal performance of embryonic development rate yields accurate prediction of embryonic age in wild reptile nests.

Temperature has a strong effect on ectotherm development rate. It is therefore possible to construct predictive models of development that rely solely on temperature, which have applications in a range of biological fields. Here, we leverage a reference series of development stages for embryos of the turtle Chelydra serpentina, which was described at a constant temperature of 20 °C. The reference series acts to map each distinct developmental stage onto embryonic age (in days) at 20 °C. By extension, an embryo taken from any given incubation environment, once staged, can be assigned an equivalent age at 20 °C. We call this concept "Equivalent Development", as it maps the development stage of an embryo incubated at a given temperature to its equivalent age at a reference temperature. In the laboratory, we used the concept of Equivalent Development to estimate development rate of embryos of C. serpentina across a series of constant temperatures. Using these estimates of development rate, we created a thermal performance curve measured in units of Equivalent Development (TPCED). We then used the TPCED to predict developmental stage of embryos in several natural turtle nests across six years. We found that 85% of the variation of development stage in natural nests could be explained. Further, we compared the predictive accuracy of the model based on the TPCED to the predictive accuracy of a degree-day model, where development is assumed to be linearly related to temperature and the amount of accumulated heat is summed over time. Information theory suggested that the model based on the TPCED better describes variation in developmental stage in wild nests than the degree-day model. We suggest the concept of Equivalent Development has several strengths and can be broadly applied. In particular, studies on temperature-dependent sex determination may be facilitated by the concept of Equivalent Development, as development age maps directly onto the developmental series of the organism, allowing critical periods of sex determination to be delineated without invasive sampling, even under fluctuating temperature.