Male-male social bonding, coalitionary support and reproductive success in wild Guinea baboons.

Male-male bonds may confer substantial fitness benefits. The adaptive value of these relationships is often attributed to coalitionary support, which aids in rank ascension and female defence, ultimately resulting in greater reproductive success. We investigated the link between male-male sociality and both coalitionary support and reproductive success in wild Guinea baboons. This species lives in a tolerant multi-level society with reproductive units comprising a male and 1-6 females at the core. Males are philopatric, form differentiated, stable and equitable affiliative relationships (strong bonds) with other males, and lack a clear rank hierarchy. Here, we analysed behavioural and paternity data for 30 males and 50 infants collected over 4 years in the Niokolo-Koba National Park, Senegal. Strongly bonded males supported each other more frequently during conflicts, but strong bonds did not promote reproductive success. Instead, males that spent less time socializing with other males were associated with a higher number of females and sired more offspring. Notably, reproductively active males still maintained bonds with other males, but adjusted their social investment in relation to life-history stage. Long-term data will be needed to test if the adaptive value of male bonding lies in longer male tenure and/or in promoting group cohesion.

Vocal convergence in a multi-level primate society: insights into the evolution of vocal learning.

The extent to which nonhuman primate vocalizations are amenable to modification through experience is relevant for understanding the substrate from which human speech evolved. We examined the vocal behaviour of Guinea baboons, Papio papio, ranging in the Niokolo Koba National Park in Senegal. Guinea baboons live in a multi-level society, with units nested within parties nested within gangs. We investigated whether the acoustic structure of grunts of 27 male baboons of two gangs varied with party/gang membership and genetic relatedness. Males in this species are philopatric, resulting in increased male relatedness within gangs and parties. Grunts of males that were members of the same social levels were more similar than those of males in different social levels (N = 351 dyads for comparison within and between gangs, and N = 169 dyads within and between parties), but the effect sizes were small. Yet, acoustic similarity did not correlate with genetic relatedness, suggesting that higher amounts of social interactions rather than genetic relatedness promote the observed vocal convergence. We consider this convergence a result of sensory-motor integration and suggest this to be an implicit form of vocal learning shared with humans, in contrast to the goal-directed and intentional explicit form of vocal learning unique to human speech acquisition.

On the evolution of baboon greeting rituals.

To balance the trade-offs of male co-residence, males living in multi-male groups may exchange ritualized greetings. Although these non-aggressive signals are widespread in the animal kingdom, the repertoire described in the genus Papio is exceptional, involving potentially harmful behaviours such as genital fondling. Such greetings are among the most striking male baboon social interactions, yet their function remains disputed. Drawing on the comprehensive analysis from our own research on wild Guinea baboons, combined with a survey of the literature into other baboon species, we review the form and function of male-male ritualized greetings and their relation to the various social systems present in this genus. These ritualized signals differ between species in their occurrence, form and function. While ritualized greetings are rare in species with the most intense contest competition, the complexity of and risk involved in greeting rituals increase with the degree of male-male tolerance and cooperation. The variety of societies found in this genus, combined with its role as a model for human socioecological evolution, sheds light on the evolution of ritualized behaviour in non-human primates and rituals in humans. This article is part of the theme issue 'Ritual renaissance: new insights into the most human of behaviours'.

Greetings in male Guinea baboons and the function of rituals in complex social groups.

Ritualized greetings, defined as exchanges of non-aggressive signals, are common among males living in multi-male groups and are thought to balance the trade-offs of male co-residence. While ritualized greetings are widespread in the animal kingdom, the behavioral repertoire described in the genus Papio is exceptional, as it involves potentially harmful behaviors such as genital fondling. Although greetings are one of the most striking male social interactions in baboons, their function is still disputed. We investigated the function of male-male ritualized greeting behavior in wild Guinea baboons. This species lives in multilevel societies where several 'units' comprising a primary male, females with young, and occasionally a secondary male form a 'party', and two to three parties form a gang. Adult males maintain affiliative relationships with preferred male partners whom they support in coalitions, regardless of kinship. We examined the social behavior of 24 adolescent and adult males (∼900 h focal observations) in the Niokolo-Koba National Park, Senegal, to test whether greetings reflect relationship quality or function to buffer tension. Greetings were ten times more frequent than aggression and twice as frequent as affiliation. Neither dyadic aggression nor tense context predicted greeting rate, discounting the buffering hypothesis. Greetings occurred almost exclusively between males of the same party, even when other parties were around. Within parties, spatially tolerant partners greeted more frequently but dyads did not greet due to proximity prior to the greeting. Although affiliation did not predict overall greeting rate, intense and potentially costly greetings were more likely between males with stronger affiliative relationships. Greetings in Guinea baboons appear to signal commitment among party members, test relationships among spatially tolerant partners, and accentuate relationship strength among highly affiliated males. Although ritualized baboon greetings lack the symbolic component of human rituals, they appear to serve similar functions, specifically to strengthen in-group affiliation and promote cooperation.

Charting the neglected West: The social system of Guinea baboons.

OBJECTIVES: Primate social systems are remarkably diverse, and thus play a central role in understanding social evolution, including the biological origin of human societies. Although baboons have been prominently featured in this context, historically little was known about the westernmost member of the genus, the Guinea baboon (Papio papio). MATERIAL AND METHODS: Here, we summarize the findings from the first years of observations at the field site CRP Simenti in the Niokolo Koba National Park in Senegal. RESULTS: Guinea baboons reveal a nested multi-level social organization, with reproductive units comprising one "primary" male, one to several females, young, and occasionally "secondary" males at the base of the society. Three to five units form "parties," which team up with other parties to form a "gang." Different gangs have largely overlapping home ranges and agonistic interactions between different parties or gangs are rare. Some but not all strongly socially bonded males are highly related, and population genetic and behavioral evidence indicate female-biased dispersal. Females play an important role in intersexual bond formation and maintenance, and female tenure length varies between a few weeks to several years. DISCUSSION: While the social organization resembles that of hamadryas baboons (P. hamadryas), the social structure differs considerably, specifically in terms of low male aggressiveness and female freedom. Despite substantial differences in social organization and social structure, the acoustic structure of Guinea baboon vocalizations does not differ substantially from that of other baboon taxa. With its multi-level organization, stable bonds between males and females, as well as a high-degree of male-male cooperation and tolerance, Guinea baboons constitute an intriguing model for reconstructing human social evolution.