ON THE MEASUREMENT OF NATURAL AND SEXUAL SELECTION: APPLICATIONS.

In this paper, we use measures of selection developed by quantitative geneticists and some new results (Arnold and Wade, 1984) to analyze multiple episodes of selection in natural populations of amphibians, reptiles, and insects. These examples show how different methods of data collection influence the potential for relating field observations to formal evolutionary theory. We adhere to the Darwinian tradition of distinguishing between natural and sexual selection (Darwin, 1859, 1871; Ghiselin, 1974). We view sexual selection as selection arising from variance in mating success and natural selection as arising from variance in other components of fitness. The justification for this formal distinction is developed by Wade (1979), Lande (1980), Wade and Arnold (1980), Arnold and Houck (1982) and Arnold (1 983 a). (We define mating success as the number of mates that bear progeny given survival of the mating organsim to sexual maturity. We do not equate mating success with mere copulatory success.) The utility of the distinction between sexual and natural selection is that the two forms of selection may often act in opposite directions on particular characters (Darwin, 1859, 1871). While we find the distinction between these two forms of selection useful, the difference is not crucial to our analysis. The essential point is that the recognition of selection episodes permits analysis of selection that may change in magnitude and direction during the life cycle. Defining Fitness Components. -The key first step in the analysis of data is to define multiplicative components of fitness so that selection can be partitioned into parts corresponding to these components or episodes of selection. Using an animal example, if the number of offspring zygotes is taken as total fitness, we can define the following components of fitness: viability (survivorship to sexual maturity), mating success (the number of mates) and fertility per mate (the average number of zygotes produced per mate). These components of fitness are defined so that their product gives total fitness. As a second example, consider the components of fitness in a plant in which yield (seeds/plant) is taken as the measure of total fitness (Primack and Antonovics, 1981). We might define the following components of fitness: number of stems per plant, average number of inflorescences per stem, average number of seed capsules per inflorescence, and average number of seeds per capsule. Again, these four fitness components are defined so that their product gives total fitness. We will need to measure each component of fitness and each character on each individual in order to partition selection into parts corresponding to the separate episodes of selection or to the separate components of fitness. Thus in the animal example, we need to measure the viability, mating success and fertility of each individual. With this accomplished we can estimate the separate forces of viability, sexual and fertility selection on each phenotypic character. In addition we can calculate the opportunities of selection corresponding to these three episodes and covariances between the different kinds of selection. In the plant example, we might begin with the intuition that larger plants have a greater yield. Using our methodology we can reword and extend this intuition. We can not only test the proposition of