Lifetime reproductive output: individual stochasticity, variance, and sensitivity analysis.

Lifetime reproductive output (LRO) determines per-generation growth rates, establishes criteria for population growth or decline, and is an important component of fitness. Empirical measurements of LRO reveal high variance among individuals. This variance may result from genuine heterogeneity in individual properties, or from individual stochasticity, the outcome of probabilistic demographic events during the life cycle. To evaluate the extent of individual stochasticity requires the calculation of the statistics of LRO from a demographic model. Mean LRO is routinely calculated (as the net reproductive rate), but the calculation of variances has only recently received attention. Here, we present a complete, exact, analytical, closed-form solution for all the moments of LRO, for age- and stage-classified populations. Previous studies have relied on simulation, iterative solutions, or closed-form analytical solutions that capture only part of the sources of variance. We also present the sensitivity and elasticity of all of the statistics of LRO to parameters defining survival, stage transitions, and (st)age-specific fertility. Selection can operate on variance in LRO only if the variance results from genetic heterogeneity. The potential opportunity for selection is quantified by Crow's index I , the ratio of the variance to the square of the mean. But variance due to individual stochasticity is only an apparent opportunity for selection. In a comparison of a range of age-classified models for human populations, we find that proportional increases in mortality have very small effects on the mean and variance of LRO, but large positive effects on I . Proportional increases in fertility increase both the mean and variance of LRO, but reduce I . For a size-classified tree population, the elasticity of both mean and variance of LRO to stage-specific mortality are negative; the elasticities to stage-specific fertility are positive.