The units of selection and measures of fitness.

This paper presents a unified account of the properties of the measures, Malthusian parameter and entropy in predicting evolutionary change in populations of macromolecules, cells and individuals. The Malthusian parameter describes the intrinsic rate of increase of the population. The entropy describes the intrinsic variability in populations: it characterizes the variability in mutation and replication rates in populations of macromolecules; the rate of decay of synchrony in populations of cells; and the degree of iteroparity in populations of individuals. The Malthusian parameter determines ultimate population numbers: under constant environmental conditions, it is the rate of increase when equilibrium conditions are attained. Entropy determines population stability: the gain in the Malthusian parameter due to small fluctuations in the life-cycle variables is determined by entropy. These properties, which are valid for populations of macromolecules, cells and individuals, show that the Malthusian parameter and entropy act as complimentary fitness indices in understanding evolutionary change in populations.