Using a behavior random permutation model to identify displacement grooming in ungulates

Behavior affects an individual's life in all aspects, e.g., enhancing fitness, leveraging predation risk, and reducing competition with conspecifics. However, the sequential distribution of behaviors received less attention and is unclear what the function of displacement behavior is. Displacement activities can be found in vertebrate species but there is no formal method to determine whether a behavior is expressed as a displaced or normal activity. Analyzing the sequential distributions of behaviors in a natural setting may allow researchers to identify unexpected distributions as a possible signature of displacement activities. In this study, we used a behavior random permutation model to detect the presence of a displacement activity in the Tibetan antelope Pantholops hodgsonii and the Tibetan gazelle Procapra picticaudata. The results showed that grooming in both ungulates tended to be accompanied with vigilance, and the frequency of grooming after vigilance was significantly higher than before vigilance. A significant positive correlation between the scan rate and grooming rate in the 2 ungulates was obtained. We suggest that grooming could sometimes be expressed as a displacement activity in ungulates. In addition to providing a general method for further research on displacement activities in a variety of animal species, this study sheds light on the importance of a spectral analysis of sequential distribution of animal behaviors. Behavior random permutation models can be used to explore the relevance between any 2 behaviors in a specific sequence, especially to identify a myriad of unexpected behaviors relative to their normal context of occurrence.