Density-dependent parameters and demographic equilibrium in open populations

Populations near their equilibrium are expected to show density-dependence through a negative feedback on at least one demographic parameter, e.g. survival and/or productivity. Nevertheless, it is not always clear which vital rate is affected the most, and even less whether this dependence holds in open populations in which immigration and emigration are also important. We assessed the relative importance of population density in the variation of local survival, recruitment, proportion of transients (emigrants) and productivity through the analysis of detailed life-histories of 4286 seabirds from a colony that reached an apparent demographic equilibrium after a period of exponential increase. We provide evidence that the role of population density and resource availability changes according to the demographic parameter considered. Estimates indicated that transients increased from 5% to 20% over the study period, suggesting an average turnover of about 1400 individuals per year. The parameters most influenced by population density alone were local survival and probability of transience. Recruitment was negatively associated with population density during the increasing phase but unexpected high values were also recorded at high population levels. These high values were explained by a combination of population size and food availability. Mean productivity varied with food availability, independently from population variations. The population density alone explained up to a third of the yearly variation of the vital rates considered, suggesting that open populations are equally influenced by stochastic and density-independent events (such as environmental perturbations) than by intrinsic (i.e. density-dependent) factors.