Social Behavior and Group Formation in Male Asian Elephants (Elephas maximus): The Effects of Age and Musth in Wild and Zoo-Housed Animals

SIMPLE SUMMARY: The conservation of wild and zoo-housed Asian elephants partly depends on developing our understanding of male elephant social behavior. Once thought to be solitary, we now know that male elephants can display rich social exchanges with both males and females. However, these interactions are expected to change with age and around the sexually active state of “musth.” We used behavioral data from wild and zoo-housed elephant populations to investigate how age, musth, and environmental factors influence how male elephants socialize and associate with group members. In Sri Lanka, only male elephants of older age (>20 years) exhibited signs of musth, but even some of the youngest zoo-housed elephants underwent musth (as young as 11 years). Additionally, we found that age and musth status predicted whether a wild male elephant associated with females, other males or was alone, as well as the number of conspecifics (males and females) in the same group with which a male was observed. Finally, rates of aggression, prosocial behavior (affiliative behaviors that promote positive social bonds between elephants), and submissive behavior exhibited by wild and zoo-housed male elephants were associated with age, musth status, the number of elephants presented, and group type (all-male or mixed sex). These results provide motivation for future studies of social behavior in male Asian elephants, as they will contribute to the reproduction and conservation of this endangered species. ABSTRACT: Asian elephants are endangered, and the long-term viability of the species depends on integrative approaches to address the sustainability of in-situ and ex-situ populations. Growing evidence shows that male elephants exhibit extensive and flexible social behavior that rivals the complexity of that of females. Male elephant sociality is expected to change dramatically around the unique sexual state of musth. However, data related to male Asian elephant sociality is lacking. Here, we conducted complementary observations in Wasgamuwa National Park, Sri Lanka, and North American zoos of male Asian elephant social behavior. Age and musth status, along with other factors, were associated with variation in social behavior and group formation of males. In wild male elephants, both musth status and age impacted elephant associations within all-male and mixed-sex groups: non-musth elephants were generally sighted less often in mixed-sex groups as they aged, while the inverse occurred with musth elephants. Musth status interacted with age to predict the number of conspecifics with which a wild male elephant associated: younger males were observed with more females during non-musth (but the opposite was true during musth), and male elephants between 20 and 30 years were observed with the highest number of male conspecifics except during musth. Finally, we found variation in aggression, prosocial behavior, and submissive behavior was influenced by intrinsic (age and musth status) and extrinsic factors (group size and type) in similar ways in both populations; prosocial behavior was most common and was influenced by the number of conspecifics present (both populations), and age, group type, and musth status (zoo population), while aggression was rare, especially among older elephants. We suggest that longitudinal studies of this threatened species will be particularly helpful to promote the reproduction and conservation of Asian elephants in in-situ and ex-situ environments.