Mating behavior, population growth, and the operational sex ratio: a periodic two-sex model approach.

We present a new approach to modeling two-sex populations, using periodic, nonlinear two-sex matrix models. The models project the population growth rate, the population structure, and any ratio of interest (e.g., operational sex ratio). The periodic formulation permits inclusion of highly seasonal behavioral events. A periodic product of the seasonal matrices describes annual population dynamics. The model is nonlinear because mating probability depends on the structure of the population. To study how the vital rates influence population growth rate, population structure, and operational sex ratio, we used sensitivity analysis of frequency-dependent nonlinear models. In nonlinear two-sex models the vital rates affect growth rate directly and also indirectly through effects on the population structure. The indirect effects can sometimes overwhelm the direct effects and are revealed only by nonlinear analysis. We find that the sensitivity of the population growth rate to female survival is negative for the emperor penguin, a species with highly seasonal breeding behavior. This result could not occur in linear models because changes in population structure have no effect on per capita reproduction. Our approach is applicable to ecological and evolutionary studies of any species in which males and females interact in a seasonal environment.