PROXIMATE AND ULTIMATE CONTROLS ON LIFE-HISTORY VARIATION: THE EVOLUTION OF LITTER SIZE IN WHITE-FOOTED MICE...

Recruitment of litter-mates of nest-box-inhabiting white-footed mice was monitored to study the evolution of litter size. The frequency distribution of litter sizes was nonsymmetrical, and the most frequent litter size was less than the optimum. This was not the result of differential parental survival, which was independent of litter size produced. Recruitment remained constant or increased slightly to a peak in litters of five young, and then dropped precipitously for larger litters. The single optimum litter size of five did not appear to have any physiological correlates. Instead, the equally low probability of successful recruitment of any young from any given litter may have given rise to a bet-hedging strategy of frequent iterated reproductions. A theoretical analysis of optimal parental investment in offspring was initiated under the assumption that optimal brood size represents a maximization of differences between age-specific costs and benefits of reproduction, both of which should be measured in constant currency of inclusive fitness. In the past, benefit has been measured by current fecundity, and cost by residual reproductive value. However, reproductive value is an appropriate estimate of inclusive fitness only for organisms in which parental investment has little effect on the subsequent survival of offspring to reproductive age. Reproductive value weighted by offspring survival and devalued by the degree of genetic relatedness defines a new currency, replacement value, which is more appropriate for evaluating the costs and benefits of parent-offspring conflict over parental investment in current as opposed to future young. Total parent-offspring conflict intensifies with increases in current brood size. For species with severe reproductive constraints, such as post-partum estrus in white-footed mice, such conflict may force parents to curtail investment in current offspring at or near parturition of subsequent litters, even if that means reducing the survival of current young.