Surplus killing by carnivores

Kruuk documented and analyzed cases where carnivores kill far more prey than they can consume, challenging the assumption that predators only take what they need. Drawing on spotted hyena and other carnivore observations, he showed that surplus killing typically occurs when prey anti-predator behavior fails, for example among confused herds at night, in storms, or with naive or trapped prey, so that the predator's normal stopping cue (prey escape) never fires. The paper framed surplus killing as a byproduct of evolved hunting motivation rather than wastefulness, and it remains the foundational reference for the phenomenon, which has since been recorded in lions, leopards, pumas, lynx and many other felids under permissive conditions.