Some non-human primates produce acoustically distinct alarm calls to different predators, such as eagles or leopards. Recipients respond to these calls as if they have seen the actual predator, which has led to the notion of functionally referential alarm calls. However, in a previous study with free-ranging putty-nosed monkeys (Cercopithecus nictitans martini), we demonstrated that callers produced two acoustically distinct alarm calls to eagle shrieks and leopard growls, but both alarm calls were given to both predators. We can think of two basic explanations for this surprising result, a methodological and theoretical one. Firstly, acoustic predator models may not always be suitable to test alarm call behaviour in primates, sometimes causing uncharacteristic behaviour. Secondly, referential alarm calling may not be a universal feature of primate alarm call systems. Considering the methodological and theoretical importance of these possibilities, we conducted a follow-up study using life-sized leopard, eagle, and human models on the same population and compared the resulting vocal responses to those given to acoustic predator models. We compared the alarm call series given to each of these predator model types and found a considerable degree of consistency suggesting that the mode of presentation did not affect anti-predator calling strategies. However, evidence for audience effects on calling behaviour was inconclusive. While it appears that predator class is reliably encoded by different call series types irrespective of the mode of presentation, observations of these same call series given in non-predatory contexts indicate that predator class is unlikely to be the relevant organising principle underlying the alarm-calling behaviour in this species. We conclude by offering an alternative, non-referential, account of the alarm-calling system exhibited by this species.
Referential communication Anti-predator Call combinations Semantic Syntax Predation Evolution of language Cognition
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We are grateful to Volker Sommer for his permission and support to conduct research on monkey vocalisations as part of the Gashaka Primate Project. We thank the Nigerian National Parks Service for providing the research permits and the Nigerian Conservation Foundation (NCF) for help with logistics. We thank Dan Azumi and Ali Tappare for their invaluable help in the field. We also thank Robert Seyfarth and three anonymous reviewers for helpful comments and criticisms. This research has been funded by research grants of the Wenner Gren Foundation, the European Science Foundation (‘Origins of Man, Language, and Languages’), the British Academy, the Leverhulme Trust, and the EC Pathfinder FP6 (‘Origins of referential communication’).
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