Predator effects on prey demography have traditionally been ascribed solely to direct killing in studies of population ecology and wildlife management. Predators also affect the prey's perception of predation risk, but this has not been thought to meaningfully affect prey demography. We isolated the effects of perceived predation risk in a free-living population of song sparrows by actively eliminating direct predation and used playbacks of predator calls and sounds to manipulate perceived risk. We found that the perception of predation risk alone reduced the number of offspring produced per year by 40%. Our results suggest that the perception of predation risk is itself powerful enough to affect wildlife population dynamics, and should thus be given greater consideration in vertebrate conservation and management.

Summary1. Predator-induced stress has been used to exemplify the concept of stress for close to a century because almost everyone can imagine the terror of fleeing for one's life from a lion or a tiger. Yet, because it has been assumed to be acute and transitory, predator-induced stress has not been much studied by either comparative physiologists or population ecologists, until relatively recently. 2. The focus in biomedical research has always been on chronic stress in humans, which most comparative physiologists would agree results from 'sustained psychological stress -linked to mere thoughts' rather than 'acute physical crises' (like surviving a predator attack) or 'chronic physical challenges' (such as a shortage of food). Population ecologists have traditionally focused solely on the acute physical crisis of surviving a direct predator attack rather than whether the risk of such an attack may have a sustained effect on other demographic processes (e.g. the birth rate). 3.Demographic experiments have now demonstrated that exposure to predators or predator cues can have sustained effects that extend to affecting birth and survival in free-living animals, and a subset of these have documented associated physiological stress effects. These and similar results have prompted some authors to speak of an 'ecology of fear', but others object that 'the cognitive and emotional aspects of avoiding predation remain unknown'. 4. Recent biomedical studies on animals in the laboratory have demonstrated that exposure to predators or predator cues can induce 'sustained psychological stress' that is directly comparable to chronic stress in humans, and this has now in fact become one of the most common stressors used in studies of the animal model of post-traumatic stress disorder (PTSD). 5. We review these recent findings and suggest ways the laboratory techniques developed to measure the 'neural circuitry of fear' could be adapted for use on free-living animals in the field, in order to: (i) test whether predator risk induces 'sustained psychological stress' in wild animals, comparable to chronic stress in humans and (ii) directly investigate 'the cognitive and emotional aspects of avoiding predation' and hence the 'ecology of fear'.

The never-ending tension between finding food and avoiding predators may be the most universal natural stressor wild animals experience. The 'chronic stress' hypothesis predicts: (i) an animal's stress profile will be a simultaneous function of food and predator pressures given the aforesaid tension; and (ii) these inseparable effects on physiology will produce inseparable effects on demography because of the resulting adverse health effects. This hypothesis was originally proposed to explain synergistic (inseparable) food and predator effects on demography in snowshoe hares (Lepus americanus). We conducted a 2 Â 2, manipulative food addition plus natural predator reduction experiment on song sparrows (Melospiza melodia) that was, to our knowledge, the first to demonstrate comparable synergistic effects in a bird: added food and lower predator pressure in combination produced an increase in annual reproductive success almost double that expected from an additive model. Here we report the predicted simultaneous food and predator effects on measures of chronic stress in the context of the same experiment: birds at unfed, high predator pressure (HPP) sites had the highest stress levels; those at either unfed or HPP sites showed intermediate levels; and fed birds at low predator pressure sites had the lowest stress levels.

The fear large carnivores inspire, independent of their direct killing of prey, may itself cause cascading effects down food webs potentially critical for conserving ecosystem function, particularly by affecting large herbivores and mesocarnivores. However, the evidence of this has been repeatedly challenged because it remains experimentally untested. Here we show that experimentally manipulating fear itself in free-living mesocarnivore (raccoon) populations using month-long playbacks of large carnivore vocalizations caused just such cascading effects, reducing mesocarnivore foraging to the benefit of the mesocarnivore's prey, which in turn affected a competitor and prey of the mesocarnivore's prey. We further report that by experimentally restoring the fear of large carnivores in our study system, where most large carnivores have been extirpated, we succeeded in reversing this mesocarnivore's impacts. We suggest that our results reinforce the need to conserve large carnivores given the significant “ecosystem service” the fear of them provides.

Apex predators such as large carnivores can have cascading, landscape‐scale impacts across wildlife communities, which could result largely from the fear they inspire, although this has yet to be experimentally demonstrated. Humans have supplanted large carnivores as apex predators in many systems, and similarly pervasive impacts may now result from fear of the human ‘super predator’. We conducted a landscape‐scale playback experiment demonstrating that the sound of humans speaking generates a landscape of fear with pervasive effects across wildlife communities. Large carnivores avoided human voices and moved more cautiously when hearing humans, while medium‐sized carnivores became more elusive and reduced foraging. Small mammals evidently benefited, increasing habitat use and foraging. Thus, just the sound of a predator can have landscape‐scale effects at multiple trophic levels. Our results indicate that many of the globally observed impacts on wildlife attributed to anthropogenic activity may be explained by fear of humans.