text: Spencer and Janzen (2014) found further support for the Charnov-Bull model by incubating painted turtles (Chrysemys picta) at different temperatures and measuring various characteristics indicative of fitness. The turtles were incubated at temperatures that produce solely males, both sexes, and solely females. Spencer and Janzen (2014) found that hatchlings from mixed-sex nests were less energy efficient and grew less than their same-sex counterparts incubated in single-sex producing temperatures. Hatchlings from single-sex producing temperatures also had higher first-year survivorship than the hatchlings from the temperature that produces both sexes. TSD may be advantageous and selected for in turtles, as embryo energy efficiency and hatchling size are optimized for each sex at single-sex incubation temperatures and are indicative of first-year survivorship. This suggests that natural selection would favor TSD, as TSD may enhance the fitness of offspring.
  An alternative hypothesis of adaptive significance was proposed by Bulmer and Bull in 1982 and supported by the work of Pen et al. (2010). They conjectured that disruptive selection produced by variation in the environment could result in an evolutionary transition from ESD to GSD. Pen et al. (2010) addresses evolutionary divergence in SDMs via natural selection on sex ratios. Studying the spotted skink, they observed that the highland population was not affected by temperature, yet, there was a negative correlation between annual temperature and cohort sex ratios in the lowlands. The highlands are colder with a higher magnitude of annual temperature fluctuation and a shorter activity season, delaying maturity, thus GSD is favored so sex ratios are not skewed. However, in the lowlands, temperatures are more constant and a longer activity season allows for favorable conditions for TSD. They concluded that this differentiation in climate causes divergent selection on regulatory elements in the sex-determining network allowing for the emergence of sex chromosomes in the highlands.