A primate grammar enabling incremental processing.

Characterizing the structure and function of animal communication systems provides insights into the cognitive and evolutionary processes shaping signal complexity. One key question is whether and how call sequences allow potential listeners to make predictions about the call-eliciting referents. Here, we investigated whether primate call sequences contained properties that enabled such predictive processing. We analyzed several years of experimentally elicited alarm responses from a West African forest primate, wild olive colobus monkeys. Using Kullback-Leibler divergence and prediction gain approaches, we identified a simple primate grammar that allowed predictions of referents from only minimal input. In particular, sequence-initial positions reliably discriminated urgent from non-urgent threats while the following positions increased the referential specificity regarding two main predators (eagles and leopards) and non-predatory disturbances (falling tree parts). Sequences often contained further calls, which may allow callers to either confirm the referent or to alter the conveyed information. We concluded that animal communication can contain features adapted for predictive, incremental processing, suggesting evolutionary roots older than language.