Richardson’s ground squirrels ( Spermophilus richardsonii (Sabine, 1822)) produce audible (ca. 8 kHz) and (or) ultrasonic (ca. 48 kHz) alarm vocalizations that warn conspecifics of impending danger. Audible calls have a larger active space than ultrasonic calls because they travel farther, are louder, and contain frequencies to which conspecific and allospecific recipients are more sensitive. In our first experiment, we presented an alarming stimulus to 103 squirrels to examine the effect of threat proximity on signal type. The ratio of ultrasonic to audible alarm calls increased with increasing distance from the stimulus. We conclude that the size of the active space influences signalling strategy and that squirrels emitting ultrasonic calls can signal conspecifics to the exclusion of distant predators. As recipients of ultrasonic calls must be close to the signaler, one context in which ultrasonic calling may be most adaptive is during natal emergence when juveniles are particularly abundant, highly vulnerable to predation, and clustered in space. In our second experiment, we broadcast ultrasonic alarm signals to emerging juveniles and found that they, like older individuals, responded to calls by increasing vigilance. We discuss the adaptive utility of multiple signalling strategies in light of our findings.
Sex differences in helping effort reveal the effect of future reproduction on cooperative behaviour in birds
