Antipredator responses of California ground squirrels to rattlesnakes and rattling sounds: the roles of sex, reproductive parity, and offspring age in assessment and decision-making rules

Abstract

California ground squirrels (Spermophilus beecheyi) and northern Pacific rattlesnakes (Crotalus viridus oreganus) have an adversarial relationship. Adults are partially protected by venom resistance and harass rattlesnakes in part to defend their more vulnerable offspring. Larger, warmer snakes are more dangerous than smaller colder snakes, and in escalated conflict squirrels could benefit from risk assessment strategies. Rattlesnakes often rattle at harassing squirrels and rattling sounds produce cues related to body size and temperature. In study 1 we played back rattling sounds from snakes that varied in dangerousness and evaluated the roles of sex and parity in squirrel risk assessment strategies. In general, squirrels tail flagged and stood bipedally more, and were slower to reapproach the playback speaker following playbacks of rattling sounds from more dangerous snakes. In comparison with males and nonmothers, mothers were most responsive to rattling sounds and more sensitive to variation in snake dangerousness. Mothers tail flagged more than males and nonmothers, and this behavior tracked variation in snake dangerousness most closely, perhaps reflecting the effects of snake size and temperature on pup vulnerability. These findings suggest that many aspects of squirrel antisnake behavior are governed by their effects on descendant kin. In study 2 we tested the effects of offspring age on mothers’ responses to live rattlesnakes and rattling sounds. According to the offspring value hypothesis, mothers should take more risks in defense of older offspring because they are more likely to survive to reproductive age. By contrast, under the offspring vulnerability hypothesis, older offspring are less vulnerable to predators and thus mothers should take fewer risks. Risk-taking, as measured by behaviors that bring the squirrel close to the snake’s strike range, was either unaffected by or negatively correlated with offspring age. Thus, our findings suggest that whereas offspring value is unimportant in squirrel antisnake behavior, offspring vulnerability may affect maternal defense. We suggest that offspring vulnerability in mammals, in comparison with birds, may play a larger role in parental defense against predators.