Predator–prey studies often assume a three trophic level system where predators forage free from any risk of predation. Since meso-predators themselves are also prospective prey, they too need to trade-off between food and safety. We applied foraging theory to study patch use and habitat selection by a meso-predator, the red fox. We present evidence that foxes use a quitting harvest rate rule when deciding whether or not to abandon a foraging patch, and experience diminishing returns when foraging from a depletable food patch. Furthermore, our data suggest that patch use decisions of red foxes are influenced not just by the availability of food, but also by their perceived risk of predation. Fox behavior was affected by moonlight, with foxes depleting food resources more thoroughly (lower giving-up density) on darker nights compared to moonlit nights. Foxes reduced risk from hyenas by being more active where and when hyena activity was low. While hyenas were least active during moon, and most active during full moon nights, the reverse was true for foxes. Foxes showed twice as much activity during new moon compared to full moon nights, suggesting different costs of predation. Interestingly, resources in patches with cues of another predator (scat of wolf) were depleted to significantly lower levels compared to patches without. Our results emphasize the need for considering risk of predation for intermediate predators, and also shows how patch use theory and experimental food patches can be used for a predator. Taken together, these results may help us better understand trophic interactions.
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Thanks to Beer Sheva Zoo, Aviv (Ranger, Holot Mashabim Nature Reserve) and the Israel Nature and National Parks Protection Authority. We would like to thank Amos Bouskila for his valuable advice at all stages of this study. We also thank Yaron Ziv and Aziz Subach for their ideas and suggestions. We thank Toby Goldberg for her assistance during fieldwork. M.Z. would like to thank Yaffa and Itay for their support and immense technical help. We would also like to thank Joel Brown, Barney Luttbeg and an anonymous reviewer for their valuable comments. The experiments comply with the current laws of Israel.
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