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Heterogeneity within a population is a pervasive challenge for studies of individual life-histories. Population-level patterns in age-specific reproductive success can be broken down into relative contributions from selective disappearance, selective appearance of individuals into the study population, and average change in performance for survivors (average ontogenetic development). In this article, we provide an exact decomposition. We apply our formula to data on the reproductive performance of a well characterized population of common terns (Sterna hirundo). We show that improvements with age over most of adult life and senescence at old ages are primarily due to a genuine change in the mean among surviving individuals rather than selective disappearance or selective appearance of individuals. Average ontogenetic development accounts for approximately 87% of the overall age-specific population change.

Original publication




Journal article


Proc Natl Acad Sci U S A

Publication Date





7841 - 7846


Age Factors, Aging, Animals, Charadriiformes, Germany, Models, Biological, Observation, Population Dynamics, Reproduction