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The fear large carnivores inspire in large ungulates has been argued to have cascading effects down food webs. However, a direct link between ungulate habitat use and their fear of large carnivores has not been experimentally tested. To fill this critical gap, we conducted a bi-factorial experiment in an African savanna. We removed shrub cover and broadcast large carnivore vocalizations (leopard, hyena, dog) or non-threatening control vocalizations in both experimentally cleared and shrubby control sites. We recorded the proactive (frequency of visitation) and reactive (fleeing or vigilance) responses of multiple prey (impala, warthog, nyala and bushbuck). Critically, we found a significant proactive–reactive interaction. Ungulates were 47% more likely to run after hearing a predator vocalization in shrubby control sites than experimental clearings, demonstrating that ungulates perceived less fear from large carnivores in open habitat (clearings). Consistent with this finding, ungulates visited clearings 2.4 times more often than shrubby control sites and visited shrubby control sites less often at night, when large carnivores are most active. Combined with results from previous experiments demonstrating that the disproportionate use of available habitats by large ungulates can alter ecosystem properties, our experiment provides critical evidence that the fear large carnivores inspire in large ungulates can cause trophic cascades.
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