Call concatenation in wild meerkats

Repertoire size, frequently determined by the number of discrete call types, has been used to assess vocal complexity in animals. However, species can also increase their communicative complexity by using graded signals or by combining individual calls. Animal call sequences can be divided into two main categories, each subdivided into two classes: repetitions, with either an unlimited or finite number of iterations of the same call type, and mixed call combinations, composed of two or more graded or discrete call types. Social contexts involve a wide range of behaviours and, unlike predation contexts, can be associated with both positive and negative emotions. Therefore, interactions linked to social contexts may place additional demands on an animal's communicative system and lead to the use of call com- binations. We systematically documented call combinations produced by wild meerkats, Suricata sur- icatta, a highly social carnivore, in social contexts in their natural habitat. We observed 12 distinct call combinations belonging to all four classes of combination, produced in all the observed behavioural contexts. Four combinations were each produced in a specific context whereas the remaining eight were produced in several contexts, albeit in different proportions. The broad use of combinations suggests that they represent a non-negligible part of meerkat social communication and that they can be used in flexible ways across various behavioural contexts. Comparison with combinations produced in predation contexts indicated that social call combinations are more varied in number of classes and structural complexity than the former, perhaps due to the greater variety of social contexts. However, in meerkats, combinations of functionally referential calls have been documented in predation but not social contexts, suggesting that both social and predation pressures may play a role in the evolution of combinatoriality in animal communication.