Neutral theory for life histories and individual variability in fitness components.

Individuals within populations can differ substantially in their life span and their lifetime reproductive success but such realized individual variation in fitness components need not reflect underlying heritable fitness differences visible to natural selection. Even so, biologists commonly argue that large differences in fitness components are likely adaptive, resulting from and driving evolution by natural selection. To examine this argument we use unique formulas to compute exactly the variance in life span and in lifetime reproductive success among individuals with identical (genotypic) vital rates (assuming a common genotype for all individuals). Such individuals have identical fitness but vary substantially in their realized individual fitness components. We show by example that our computed variances and corresponding simulated distribution of fitness components match those observed in real populations. Of course, (genotypic) vital rates in real populations are expected to differ by small but evolutionarily important amounts among genotypes, but we show that such differences only modestly increase variances in fitness components. We conclude that observed differences in fitness components may likely be evolutionarily neutral, at least to the extent that they are indistinguishable from distributions generated by neutral processes. Important consequences of large neutral variation are the following: Heritabilities for fitness components are likely to be small (which is in fact the case), small selective differences in life histories will be hard to measure, and the effects of random drift will be amplified in natural populations by the large variances among individuals.