Costly calling: Marmots who alarm call at higher rates are less likely to survive the summer and live shorter lives.

Emitting alarm calls may be costly, but few studies have asked whether calling increases a caller's risk of predation and survival. Since observing animals calling and being killed is relatively rare, we capitalized on over 24,000 h of observations of marmot colonies and asked whether variation in the rate that yellow-bellied marmots (Marmota flaviventer) alarm called was associated with the probability of summer mortality, a proxy for predation. Using a generalized mixed model that controlled for factors that influenced the likelihood of survival, we found that marmots who called at higher rates were substantially more likely to die over the summer. Because virtually all summer mortality is due to predation, these results suggest that calling is indeed costly for marmots. Additionally, the results from a Cox survival analysis showed that marmots that called more lived significantly shorter lives. Prior studies have shown that marmots reduce the risk by emitting calls only when close to their burrows, but this newly quantified survival cost suggests a constraint on eliminating risks. Quantifying the cost of alarm calling using a similar approach in other systems will help us better understand its true costs, which is an essential value for theoretical models of calling and social behavior.