Testing domains of danger in the selfish herd: sparrowhawks target widely spaced redshanks in flocks

The main tenet of Hamilton's 'selfish herd theory' for the evolution of group living is that individual risk of being killed upon attack by a predator is greater when relatively far from conspecifics. Here we examine the role of spacing using video analysis of encounters between redshanks, Tringa totanus, in flocks on saltmarsh, and sparrowhawks, Accipiter nisus, surprise hunting from adjacent woodland. Targeted redshanks were 35% (approx. 5 body lengths) more widely spaced than their nearest non-targeted neighbours, controlling for proximity to the hawk. Although targeted redshanks were also twice as slow to escape, the effect dropped out of a model containing spacing, which alone accounted for twice as much variation as escape delay. Redshanks were more tightly spaced on the riskiest side of the flock, suggesting they attempted to compensate for the greater risk, while birds on the edges of flocks were more widely spaced than those in the centre. Our analysis controls for most of the confounding effects associated with the edge-centre comparisons that are normally used in similar studies and provides strong support for spacing-dependent differential predation risk in the wild. In general, we suggest that positive selection for tight spacing when prey are stationary is largely due to domains of danger, but that this also leads to positive selection when prey are mobile because of predator confusion.