New evidence from observations of progressions of mandrills (Mandrillus sphinx): a multilevel or non-nested society?

African papionins are well known for the diversity of their social systems, ranging from multilevel societies based on one-male-multifemale units (OMUs) to non-nested societies. However, the characteristics of Mandrillus societies are still unclear due to difficult observational conditions in the dense forests of central Africa. To elucidate the characteristics of mandrill societies and their social systems, I analysed the age-sex compositions, behaviours, and progression patterns of their horde/subgroups using videos of them crossing open places. The progressions were very cohesive, and the very large aggregations (169-442 individuals) had only 3-6 adult males (1.4-1.8 % of all individuals) and 11-32 subadult males (6.5-7.2 %). No herding behaviours were observed in the males, and most of the small clusters within the progressions were not analogous to the OMUs of a multilevel society but instead consisted of only adult females and immatures. The progressions of alert mandrills showed patterns similar to those observed in a non-nested social system: females with dependent infants were concentrated toward the rear and adult and subadult males toward the front. These results suggest that cohesive aggregations and a female-biased sex ratio are common characteristics of mandrill species. Mandrills may form female-bonded and non-nested societies, although their fission-fusion dynamics may be different from those typical of savannah baboons.