Controlling for behavioural state reveals social dynamics among male African elephants, Loxodonta africana

The drivers of social affiliation may vary over time as individuals change their goals with respect to changing environments or physical condition. Studies of companion preference rarely consider shifts in motivational state, despite the potential importance of such shifts in structuring association and population processes. Ignoring state dependence in social behaviour may weaken the ability to recognize social properties and identify their underlying drivers. Modifying established approaches, we apply a state-specific analysis to investigate social properties in male African elephants, which are thought to be weakly social. Specifically, we delineate associations during distinct sexually active and inactive periods and quantify common social metrics (network size, density, betweenness and the number and age of preferred companions) to examine how sexual states may relate to male elephant social relationships. We found that state-dependent association index values were higher and quantitative definitions of preferred companions were more conservative than those derived when sexual state was not taken into account. Preferred companions tended to be closer in age among sexually inactive dyads relative to sexually active dyads, indicating that bulls seek out age-mates when sexually inactive. Networks were larger and denser when sexually inactive. By accounting for dynamic behaviour in social systems, this study demonstrates that male African elephants show more social preference than had been previously thought.