Animals are under strong selective pressures to make correct decisions when attempting to escape an approaching predator, and not surprisingly many studies have shown that animals adjust their flight initiation behavior in response to risk. However, we have a poor understanding of animals' capability to select an appropriate flight trajectory. We investigated whether eastern gray squirrels would adjust their flight trajectory based on the relative locations of the squirrel, the approaching threat, and potential refuges. We used a person running toward a focal squirrel (N = 122) as the threat and considered the three trees nearest the squirrel and taller than 8 m to be potential refuges. Squirrels were strongly affected by the angle (θ) formed by the locations of person, squirrel, and the three nearest trees. A squirrel was less likely to run to the nearest tree (Tree 1) when θ 1 was relatively acute, but also less likely to run to Tree 1 when θ 2 was obtuse, making Tree 2 a more attractive refuge. A squirrel was more likely to run to Tree 1 if it was close and if Tree 2 was relatively far. Subtle differences in the effects of θ 1 vs. θ 2 on squirrel refuge choice support the idea that squirrels prefer a nearby refuge. Squirrels were more likely to select Trees 2 and 3 rather than Tree 1 only when θ 2 was obtuse (105 •). In contrast, most squirrels chose to run to Tree 1 when θ 1 was >65 • ; thus squirrels were more likely to choose Tree 1 even when doing so required running at least partly toward the approaching threat. The decisions made by focal squirrels provide evidence that this species' assessment of risk is highly nuanced. A great deal of variation has been reported in responses to predators within species. While part of the variation may be due to strategic unpredictability on the part of the prey, part of it may also be due to differences in flight trajectory and refuge preferences that have not been well-studied.
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