Contextual and Combinatorial Structure in Sperm Whale Vocalisations
Sperm whales (Physeter macrocephalus) are long-lived and highly social mammals that engage in complex group behaviours, including navigation, foraging, and child-rearing. During these behaviours, sperm whales communicate primarily using sequences of short bursts of clicks with varying inter-click intervals, known as codas. Past research has identified around 150 discrete coda types globally, with 21 in the Caribbean. A subset of these have been shown to encode information about caller and clan identity. However, almost everything else about the sperm whale communication system, including basic questions about its structure and information-carrying capacity, remains unknown. In this study, we show that codas exhibit contextual and combinatorial structure with key similarities to aspects of human language and other primate communication systems. First, we report previously undescribed variations in coda structure that are sensitive to the conversational context in which they occur. We call these rubato and ornamentation, by analogy to musical terminology. These variations are systematically controlled and imitated across individual whales. Second, we show that coda types are not defined by arbitrary sequences of inter-click intervals, but instead form a combinatorial coding system in which rubato and ornamentation combine with two categorical, context-independent features that we call rhythm and tempo to give rise to a large inventory of distinguishable codas. In a dataset of 8,719 codas from the sperm whales of the Eastern Caribbean clan, this sperm whale phonetic alphabet makes it possible to systematically explain observed variability in coda structure. Sperm whale vocalisations are more expressive and structured than previously believed, and are built from a repertoire comprising nearly an order of magnitude more distinguishable codas. These results show contextsensitive and combinatorial vocalisation systems extend beyond humans, and can appear in an organism with a divergent evolutionary lineage and vocal apparatus.