Combinatorics and complexity of chimpanzee (Pan troglodytes) facial signals

There have been shifts toward more systematic and standardized methods for studying non-human primate facial signals, thanks to advancements like animalFACS. Additionally, there have been calls to better integrate the study of both facial and gestural communication in terms of theory and methodology. However, few studies have taken this important integrative step. By doing so, researchers could gain greater insight into how the physical flexibility of facial signals affects social flexibility. Our study combines both approaches to examine the relationship between the flexibility of physical form and the social function of chimpanzee facial "gestures". We used chimpFACS along with established gestural ethograms that provide insights into four key gesture properties and their associated variables documented in chimpanzee gestures. We specifically investigated how the combinatorics (i.e., the different combinations of facial muscle movements) and complexity (measured by the number of discrete facial muscle movements) of chimpanzee facial signals varied based on: (1) how many gesture variables they exhibit; (2) the presence of a specific goal; and (3) the context in which they were produced. Our findings indicate that facial signals produced with vocalizations exhibit fewer gesture variables, rarely align with specific goals, and exhibit reduced contextual flexibility. Furthermore, facial signals that include additional visual movements (such as those of the head) and other visual signals (like manual gestures) exhibit more gestural variables, are frequently aligned with specific goals, and exhibit greater contextual flexibility. Finally, we discovered that facial signals become more morphologically complex when they exhibit a greater number of gesture variables. Our findings indicate that facial "gesturing" significantly enhanced the facial signaling repertoire of chimpanzees, offering insights into the evolution of complex communication systems like human language.