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A major question in ecology is how age-specific variation in demographic parameters influences population dynamics. Based on long-term studies of growing populations of birds and mammals, we analyze population dynamics by using fluctuations in the total reproductive value of the population. This enables us to account for random fluctuations in age distribution. The influence of demographic and environmental stochasticity on the population dynamics of a species decreased with generation time. Variation in age-specific contributions to total reproductive value and to stochastic components of population dynamics was correlated with the position of the species along the slow-fast continuum of life-history variation. Younger age classes relative to the generation time accounted for larger contributions to the total reproductive value and to demographic stochasticity in "slow" than in "fast" species, in which many age classes contributed more equally. In contrast, fluctuations in population growth rate attributable to stochastic environmental variation involved a larger proportion of all age classes independent of life history. Thus, changes in population growth rates can be surprisingly well explained by basic species-specific life-history characteristics.

The dynamics of an age-structured population in a fluctuating environment is determined by the stochastic individual contributions from annual survival and fecundity to the total reproductive value of the population the next year. All parameters required to describe the population dynamics are simple properties of the distribution of these individual demographic contributions, which we call individual reproductive value. The asymptotic population growth rate in the average environment and the demographic and environmental variances are respectively the mean individual reproductive value over individuals through time and the variance within and between years. Our approach leads to an intuitive understanding of demographic and environmental variances in age-structured populations and their decomposition into additive age-specific components due to survival and reproduction. We show how to apply this approach to estimate the demographic and environmental variances and their components. The estimates are based on yearly random samples of individual vital rates and require no information about the total population size.

The relative importance of life-history variables to population growth rate (lambda) has substantial consequences for the study of life-history evolution and for the dynamics of biological populations. Using life-history data for 142 natural populations of mammals, we estimated the elasticity of lambda to changes in age at maturity (alpha), age at last reproduction (omega), juvenile survival (Pj), adult survival (Pa), and fertility (F). Elasticities were then used to quantify the relative importance of alpha, omega, Pj, Pa, and F to lambda and to test theoretical predictions regarding the relative influence on lambda of changes in life-history variables. Neither alpha nor any other single life-history variable had the largest relative influence on lambda in the majority of the populations, and this pattern did not change substantially when effects of phylogeny and body size were statistically removed. Empirical support for theoretical predictions was poor at best. However, analyses of elasticities on the basis of the magnitude (F) and onset (alpha) of reproduction revealed that alpha, followed by F, had the largest relative influence on lambda in populations characterized by early maturity and high reproductive rates, or when F/alpha > 0.60. When maturity was delayed and reproductive rates were low, or when F/alpha < 0.15, survival rates were overwhelmingly most influential, and reproductive parameters (alpha and F) had little relative influence on lambda. Population dynamic consequences of likely responses of biological populations to perturbations in life-history variables are examined, and predictions are made regarding the numerical dynamics of age-structured populations on the basis of values of the F/alpha ratio.

Environmental factors influence the dynamics and regulation of biological populations through their influences on demographic variables, but demographic mechanisms of population regulation have received little attention. We investigated the demographic basis of regulation of Columbian ground squirrel (Spermophilus columbianus) populations under natural and experimentally food-supplemented conditions. Food supplementation caused substantial increases in population density, and population densities returned to pretreatment levels when the supplementation ended. Control (untreated) populations remained relatively stable throughout the study period (1981-1986). Because food resources regulated the size of the ground squirrel populations, we used life-table response experiment (LTRE) analyses to examine the demographic basis of changes in population growth rate and thus also demographic influences on population regulation. LTRE analyses of two food-manipulated populations revealed that changes in age at maturity and fertility rate of females generally made the largest contributions to observed changes in population growth rate. Thus, our results suggested that abundance of food resources regulated the size of our study populations through the effects of food resources on age at maturity and fertility rates. Our results also indicated that different demographic mechanisms can underlie population regulation under different environmental conditions, because lower juvenile survival substantially contributed to population decline, but in only one of the populations. Demographic analyses of experimental data, such as those presented here, offer a rigorous and unambiguous means to elucidate the demographic basis of population regulation and to help identify environmental factors that underlie dynamics and regulation of biological populations.

Columbian ground squirrels (Spermophilus columbianus) exhibit an elevational cline in several life-history traits and in body mass. Body mass is a trait that might interact with the elevational changes in life histories, but it could reflect either or both of two underlying factors: structural size and physiological condition. I studied these factors in several populations of ground squirrels. Structural size was estimated from first principal component scores in an analysis of postcranial bone lengths. Physiological condition was inferred from differences in body mass that were independent of structural size. Field body mass and structural size of yearlings decreased with increasing elevation, which indicates slower growth at higher elevations. For adults, physiological condition was not measured directly, but evidence suggested that it was reflected by changes in body mass: body mass declined with increasing elevation but structural size did not, residuals of body mass on structural size showed the same patterns of change as body mass, most of the variation in body mass was orthogonal to a "general size factor" in principal components analyses, and significant changes in body mass of individuals could be stimulated by experimental supplementation of food. Life-history traits and physiological condition covaried along the elevational cline and were phenotypically plastic, but the structural size of adult ground squirrels was not an extremely plastic trait.