Alarm signals are a common feature of social species and are thought to be a means of alerting others to an external threat (e.g., Gill & Bierema, 2013;Kaplan, 2014;Townsend & Manser, 2013). Among cooperatively breeding mammals and birds, individuals may give an alarm call in the presence of natural enemies, thereby allowing others to take defensive action, ranging from hiding to mobbing the intruder (Caro, 2005). The different responses to alarm calls indicate that they can convey specific information about the nature of the threat, including the kind and proximity of the predator, allowing the receivers to respond appropriately. For example, the alarm calls of vervet monkeys Chlorocebus pygerythrus differ according to the threat: avian predators elicit "eagle" calls and receivers then hide in bushes, while receivers move into trees when they hear "leopard" calls (Seyfarth et al., 1980). Thus, alarm calls are thought to have "functional reference" (Evans, 1997;Macedonia & Evans, 1993), where the receivers react to the alarm calls similarly to how they would react to the threat itself (

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