Behavioral syndromes can reduce population density: behavior and demographic heterogeneity

Behavioral syndromes are widely recognized as important for ecology and evolution, but most predictions about ecological impacts are based on conceptual models and are therefore imprecise. Borrowing insights from the theory of demographic heterogeneity, we derived insights about the population-dynamic effects of behavioral syndromes. If some individuals are consistently more aggressive than others, not just in interspecific contests, but also in foraging, mating, and anti-predator behavior, then population dynamics could be affected by the resulting heterogeneity in demographic rates. We modeled a population with a boldness-aggressiveness syndrome (with the individual's trait constant through life), showing that the mortality cost of boldness causes aggressive individuals to die earlier, on average, than their non-aggressive siblings. The equilibrium frequency of the aggressive type is strongly affected by the mortality cost of boldness, but not directly by the reproductive benefit of aggressiveness. Introducing aggressive types into a homogeneous non-aggressive population increases the average per-capita mortality rate at equilibrium; under many conditions, this reduces the equilibrium density. One such condition is that the reproductive benefit of aggression is frequency dependent and the population has evolved to equalize the expected fitness of the two types. Finally, if the intensity of aggressiveness can evolve, then the population is likely to evolve to an evolutionarily stable trait value under biologically reasonable assumptions. This analysis shows how a formal model can predict both how a syndrome affects population dynamics and how the population processes constrain evolution of the trait; we suggest some concrete predictions.