Drivers of Diversification in Individual Life Courses.

Heterogeneity in life courses among individuals of a population influences the speed of adaptive evolutionary processes, but it is less clear how biotic and abiotic environmental fluctuations influence such heterogeneity. We investigate principal drivers of variability in sequence of stages during an individual's life in a stage-structured population. We quantify heterogeneity by measuring population entropy of a Markov chain, which computes the rate of diversification of individual life courses. Using individual data of a primate population, we show that density regulates the stage composition of the population but that its entropy and the generating moments of heterogeneity are independent of density. This lack of influence of density on heterogeneity is due to neither low year-to-year variation in entropy nor differences in survival among stages but is rather due to differences in stage transitions. Our analysis thus shows that well-known classical ecological selective forces, such as density regulation, are not linked to potential selective forces governing heterogeneity through underlying stage dynamics. Despite evolution acting heavily on individual variability in fitness components, our understanding is poor whether observed heterogeneity is adaptive and how it evolves and is maintained. Our analysis illustrates how entropy represents a more integrated measure of diversity compared to the population structural composition, giving us new insights about the underlying drivers of individual heterogeneity within populations and potential evolutionary mechanisms.