Haldane's Principle

The total mutation rate corresponds to a fertility-weighted lifetable entropy expression rather than the familiar total fitness loss. The non-linear invariant differs from the linear invariant because demographic selection makes advanced-age survivors genetically selected. The paper derives a non-standard generalization of Haldane's Principle in the non-linear point-mass model. When equilibrium exists, aggregate population hazard does not depend on mutational effect sizes. Changing mutational effect size changes expected allele frequency but leaves aggregate hazard determined by mutation rates, fertility, and aggregate survivorship.