The proposed equivalence of maximizing reproductive value and fitness is examined for two model life histories. In the first instance, it is assumed that offspring are fledged before the start of the next breeding season. In this case the proposed equivalence is verified. In the second model, parents care for their progeny for more than 1 year. In this case the optimal reproductive expenditure at a particular age is shown to depend on both current reproductive value and the diminution in survival rates of previously conceived young still dependent on parental protection.Recently, there has been considerable theoretical interest in the evolution of life history phenomena (1-8). The problem, first stated explicitly by Gadgil and Bossert (4), is as follows. Consider an organism with limited (i.e., finite) resources at its disposal. Further suppose that there exists heritable variation with regard to the manner in which these resources are allocated. Then, one wishes to determine the schedule of allocation-to growth, to reproduction, and to maintenance-that maximizes the individual's contribution to subsequent generations.In attempting to answer this question, several authors (5, 6, 8, 9) have proposed that the allocation schedule in question is that which maximizes reproductive value (10) at all ages. More recently, however, this view has been questioned (1, 2). In particular, it has been suggested that maximizing fitness and reproductive value and fitness are equivalent only under the restrictive circumstance that trade-offs between current fecundity and subsequent survival and growth are confined to operate within (and not between) age classes. The purpose of the present paper is to examine this question in detail. Specifically, it is argued that maximizing fitness and reproductive value is indeed equivalent, provided that increased expenditures for reproduction at age x do not adversely affect fecundity and survivorship at previous ages y (y < x). This assumption would appear to be met in nearly all cases, save those in which there is extended parental care. In such instances, offspring as yet unfledged can suffer increased mortality if the parents breed a second time and turn their attention to the more recent litter. Mathematically, this is equivalent to reducing the parents' fecundity at earlier ages and, in such cases, natural selection can be expected to maximize reproductive value plus a weighted sum of prior years' reproduction. An analysis of this case is given.
ANALYSISPreliminaries. Consider an organism with the life cycle depicted in Fig. 1. The population is censused once a year during the breeding season, nx(t) being the number of xyear-old individuals alive in year t. Let mx be the number of offspring produced by an x-year-old individual. Of these, cx survive to be counted as young of the year (yearlings) the fol- Each of these produces mi offspring of which ci survive to the breeding season in their first year of life (zeroth age class). The product cimi = Bi is termed the effective fecun...