Simulating the effects of wolf-elk population dynamics on resource flow to scavengers

The reintroduction of gray wolves to Yellowstone National Park (YNP) provides a natural experiment regarding the effects of top predators on scavenger species. Fieldwork on the Northern Range of Yellowstone indicates that wolves facilitate carrion acqui-sition by scavengers, but it is unclear whether this represents a transient or permanent effect of wolf reintroduction. Here we present a wolf-elk model with human elk harvest and use it to investigate the long-term consequences of predator–prey dynamics and hunting on resource flow to scavengers. Our model shows that while wolves reduce the total amount of carrion, they stabilize car-rion abundance by reducing temporal variation in the quantity of carrion and extending the period over which carrion is available. Specifically, the availability of carrion is shifted from reliance on winter severity and elk density to dependence on the strength of wolf predation. Though wolves reduce the overall abundance of carrion by lowering the elk population, this reduction is partially offset by increases in the productivity of an elk population invigorated by removal of the weakest individuals. The result of this is higher carrion production per elk in the presence of wolves. In addition, this yields an ecological explanation for the phenomena that predators increase the robustness of their prey: namely that by reducing the effect of density-dependent resource competition among elk, those that remain, even some of the older animals, are better fed and healthier as a result. Our model also suggests that human hunting has no effect on the distribution of carrion across the year but is crucial in determining the long-term abundance of carrion because of the effect of hunting on elk population levels. By reducing the proportion of cows in the annual hunt, which hav