Variance and variability, uncovering an underappreciated component of reproductive isolation

Estimating the fitness of line crosses has been a key element in studies of inbreeding depression, hybridization, and speciation. Fitness values are typically compared using differences in the arithmetic mean of a fitness component between types of crosses. One aspect of fitness that is often overlooked is variance in offspring fitness over time. In the majority of studies, ignoring this aspect of fitness is unavoidable because it is impossible to estimate variance in offspring fitness over long time periods. Here, I describe a method of estimating variance in offspring fitness by substituting spatial variation for temporal variation and provide an empirical example. The method is based on Levene's test of homogeneity of variances. It is implemented by quantifying differences in residual variation among cross types. In a previous study, I performed crosses between populations of the annual plant Diodia teres and quantified hybrid fitness. In this study, another component of isolation and heterosis was revealed when considering variance in offspring fitness. When taking into account variance in offspring fitness using geometric mean fitness as the measure of performance, hybrids between populations from different habitats showed less heterosis than when calculating fitness based on arithmetic mean. This study demonstrates that variance in offspring fitness can be an important aspect of fitness that should be measured more frequently.