A general, resource-based explanation for density dependence in populations of large herbivores

The discipline of ecology seeks to understand how ecosystems, communities, and populations are regulated. A ubiquitous mechanism of population regulation of consumers is that capturing energy and nutrients in sufficient quantities for survival and reproduction becomes more difficult as population density increases. Extensive evidence has revealed that populations of large herbivores are often regulated by density dependence, defined as the reduction in the per-capita population growth rate that occurs as populations grow large. Diminished body mass of individuals has been repeatedly observed in high-density populations, implicating compromised nutrition as the primary cause of density dependence. However, there is no general explanation for why these nutritional deficiencies occur. Recent work demonstrated that reduced food intake rates resulting from the functional response of herbivores to depleted plant biomass does not provide a sensible explanation for density dependence because rates of food intake of herbivores are often insensitive to changes in plant biomass. A new model of feedbacks from plant biomass to herbivores shows how reduced nutrition of herbivores can result from increased dilution of nutrients in the plant tissue they consume as populations grow, even when their rate of consumption of plants remains constant. The model contains parameters that can be scaled to body mass, allowing unusually general predictions. The model shows that convex, concave, and linear relationships between the per-capita growth rate and population density can arise from the effects of depletion of plant biomass by herbivore foraging. The model is the first to explicitly include spatial variance in the nutritional quality of plants as a general driver of herbivore population dynamics. I show how regulation of herbivore abundance by plant nutrients can occur, even when a large fraction of the consumable plant biomass remains uneaten, providing a simple, mechanistic explanation for bottom-up control of population dynamics of primary consumers in a “green world.”