Up, up, and away: relative importance of horizontal and vertical escape from predators for survival and senescence.

Animals fleeing a potential predator can escape horizontally or vertically, although vertical flight is more expensive than horizontal flight. The ability to escape in three dimensions by flying animals has been hypothesized to result in greater survival and eventually slower senescence than in animals only fleeing in two dimensions. In a comparative study of flight initiation distance in 69 species of birds when approached by a human, I found that the amount of variance explained by flight initiation distance was more than four times as large for the horizontal than the vertical component of perch height when taking flight. The slope of the relationship between horizontal distance and flight initiation distance (horizontal slope) increased with increasing body mass across species, whereas the slope of the relationship between vertical distance and flight initiation distance (vertical slope) decreased with increasing body mass. Therefore, there was a negative relationship between horizontal and vertical slope, although this negative relationship was significantly less steep than expected for a perfect trade-off. The horizontal slope decreased with increasing density of the habitat from grassland over shrub to forest, whereas that was not the case for the vertical slope. Adult survival rate increased and rate of senescence (longevity adjusted for survival rate, body mass and sampling effort) decreased with increasing vertical, but not with horizontal slope, consistent with the prediction that vertical escape indeed provides a means of reducing the impact of predation.