Density Dependence, Evolutionary Optimization, and the Diversification of Avian Life Histories

Abstract

Abstract Although we have learned much about avian life histories during the 50 years since the seminal publications of David Lack, Alexander Skutch, and Reginald Moreau, we still do not have adequate explanations for some of the basic patterns of variation in life-history traits among birds. In part, this reflects two consequences of the predominance of evolutionary ecology thinking during the past three decades. First, by blurring the distinction between life-history traits and life-table variables, we have tended to divorce life histories from their environmental context, which forms the link between the life history and the life table. Second, by emphasizing constrained evolutionary responses to selective factors, we have set aside alternative explanations for observed correlations among life-history traits and life-table variables. Density-dependent feedback and independent evolutionary response to correlated aspects of the environment also may link traits through different mechanisms. Additionally, in some cases we have failed to evaluate quantitatively ideas that are compelling qualitatively, ignored or explained away relevant empirical data, and neglected logical implications of certain compelling ideas. Comparative analysis of avian life histories shows that species are distributed along a dominant slow-fast axis. Furthermore, among birds, annual reproductive rate and adult mortality are directly proportional to each other, requiring that pre-reproductive survival is approximately constant. This further implies that age at maturity increases dramatically with increasing adult survival rate. The significance of these correlations is obscure, particularly because survival and reproductive rates at each age include the effects of many life-history traits. For example, reproductive rate is determined by clutch size, nesting success, season length, and nest-cycle length, each of which represents the outcome of many different interactions of an individual's life-history traits with its environment. Resolution of the most basic issues raised by patterns of life histories clearly will require innovative empirical, modeling, and experimental approaches. However, the most fundamental change required at this time is a broadening of the evolutionary ecology paradigm to include a variety of alternative mechanisms for generating patterns of life-history variation.