More than a simple fixed action pattern: Yawning in drills

In the last decade, increasing attention has been devoted to exploring some aspects of yawning in non-human animals. With their chin red mark, bony paranasal swellings, male large brains and long canines, drills (Mandrillus leucophaeus) offer a robust model for testing hypotheses on the phenomenon. We identified two yawn variants (covered, YCT and uncovered teeth, YUCT) which differ in terms of recruitment of muscular action units (AUs). We tested the effects of several variables (sex, dominance rank, context) on the duration of the yawn and the probability of YCT or YUCT occurrence. We found that males performed longer and more YUCT than females. These findings support the Brain Cooling Hypothesis suggesting that those species showing large brains tend to display larger and longer yawns. We also tested the State Changing Hypothesis predicting the presence of a temporal association of yawning and ongoing behavioral transitions. A sequential analysis revealed that after 30 s following a yawn, drills were significantly more likely to change their behavioral state. Through the observation of yawning, conspecifics might gain knowledge of impending state changes. Seeing other’s yawns increased the probability of a similar response in the observers, thus suggesting the presence of yawn contagion in drills. Although the dataset needs to be expanded, our findings indicate that yawning is variable in drills, it can be associated with subjects’ state changes, and the imminent shifts can be perceived/processed by conspecifics. Graphical Abstract: (Figure presented.)