Social primates spend a significant proportion of their time exchanging grooming with their group companions. Although grooming is mainly exchanged in kind, given its hygienic and tension-reducing functions, it is still debated whether grooming also provides some social benefits, such as preferential access to resources (e.g., food or mating partners). In this study we analyzed grooming distribution among wild female Japanese macaques living in two groups on Yakushima. We tested the tendency of monkeys to reciprocate the amount of grooming received, and to direct their grooming up the hierarchy. Then we analyzed the relation of grooming to three of its possible benefits: reduced aggression, increased tolerance over food, and agonistic support against a male aggressor. The data were analyzed by means of row-wise matrix correlations. Grooming was highly reciprocated (i.e., exchanged in kind) and directed up the hierarchy in both the study groups. No significant relationship was found between grooming and aggression. Conversely, grooming favored tolerance over food, since it was positively correlated with presence on the same food patch, close proximity, and close approaches (both within 1 m) during feeding. Grooming was also positively related to agonistic support against adult males, although this relationship became nonsignificant when we controlled for kinship. Although these results are not definitive, they suggest that monkeys may derive various social benefits from grooming. This conclusion is supported by the fact that in various primate species animals tend to prefer high-ranking individuals as grooming partners.
Cords and Aureli (2000) proposed that relationship quality can be described by three components, i.e., value, security, and compatibility, based on the benefits social partners receive from their relationship and on how likely it is for them to maintain a stable friendly relationship over time. We aimed to examine whether this 3-components structure can describe social relationships and to investigate the degree of asymmetry of social relationships. We used the Japanese macaque as our model species, as group members may form strong social relationships with one another. We collected 359.2 h of focal samples, on 21 monkeys, and ran a principal component analysis (PCA) on dyadic scores for 9 variables measuring social relationships, e.g., grooming or aggression. This PCA gave 3 components. However, a second PCA run on scores per individual, i.e. for each variable and dyad we obtained 1 score for the actor and 1 for the receiver of behavior, gave a 5-component structure. We interpreted this result as evidence of the asymmetric nature of social relationships, in terms of the frequency and type of behavior exchanged between social partners, in the Japanese macaque. This interpretation was confirmed by the low correlation coefficients between behaviors given and received within a dyad. Overall these results provide mixed support for the 3-components structure predicted by Cords and Aureli (2000). They also show that social relationships may be largely asymmetric and that this asymmetry should be considered in studies on social behavior.
In many species interactions among group are often characterized by agonistic behaviour. Although animals may participate in inter-group encounters in different ways, depending on their energetic requirements, reproductive tactics, and/or developmental stage, the proximate causes affecting an animal's participation in inter-group encounters are still poorly understood. Indeed, many studies have analysed the behaviour of males and females during inter-group encounters without considering the importance of additional factors (e.g. rank). This study focuses on wild non-provisioned Japanese macaques (Macaca fuscata yakui) living on Yakushima Island, Japan. It aims to determine how monkeys of different sex, age, and rank behave during inter-group encounters and it discusses the implications and consequences of their behaviour on group composition and male dispersal. Males participated significantly more than females in inter-group encounters, by displaying more aggressive or affiliative behaviour. High-ranking and/or adult males were more aggressive than lowranking and/or subadult males during encounters occurring in the mating season and they also showed more herding behaviour. This trend was not found in inter-group encounters occurring during the non-mating season. Finally, males which then emigrated to new groups were low-ranking and/or subadult individuals. Those males displayed more affiliative behaviour towards foreign males than males which did emigrate. These data indicate that in non-territorial species with male dominance over female and high competition for mating partners males play an active, and often aggressive, role during inter-group encounter while female participation is scarce. Factors such as age, rank and period of the year (in seasonally breeding species) have to be taken into considerations when analysing interactions between groups and their effects on group composition and social behaviour.
In various social species, animals have been observed to share friendly relationships with some group members and to resolve conflicts through reconciliation, the exchange of affiliative behaviour soon after a conflict that functions to restore the relationship between the former opponents. The valuable relationship hypothesis predicts that reconciliation should be observed more often after conflicts between friends. Friendly relationships can be described by three dimensions (i.e. value, security and compatibility); however, research into the relative importance of these dimensions for the occurrence of reconciliation is sparse. Moreover, reconciliation may depend on factors other than the social relationship between opponents including, for example, their social status or the context of the conflict. Our study aimed at analysing which factors are important determinants of reconciliation and at testing the valuable relationship hypothesis, by analysing the relative effects of relationship value, security and compatibility on the occurrence and timing of reconciliation. We collected data on two troops of wild Japanese macaques living on Yakushima Island, Japan, and selected the best predicting variables of reconciliation using linear mixed models. Our results show that reconciliation occurs more frequently, and earlier, after conflicts between opponents who exchange a higher percentage of grooming. Two additional variables related to relationship security and value were selected in the best models: frequency of aggression and of approaches resulting in tolerated co‐feeding. Among the variables not related to relationship quality, distance between opponents at the end of the conflict, kinship, sex of the opponents and context of conflict (i.e. during feeding or social time) were included in our models. Our findings support the valuable relationship hypothesis and, in particular, highlight that the fitness‐related benefits of social relationships (i.e. the relationship value) are important determinants of the evolution of friendly relationships and reconciliation.
Reconciliation was first described more than 20 years ago. Since then, it has been observed in many mammals (mainly primates) but data on postconflict behavior among males are still scarce because they usually aggressively compete for mating partners, rarely maintain amicable relationships with one another. Accordingly, reconciliation is expected to occur at low rates. Although this is true for Japanese macaque males, the subspecies on Yakushima Island (Macaca fuscata yakui) seems to represent an exception as grooming among males occurs often. We analyzed postconflict behavior among them and discuss the possible factors that may favor the occurrence of grooming and reconciliation. Selective attraction between former opponents-reconciliationoccurred soon after conflicts. Consolation-affiliative interactions between a focal animal and group members other than the former opponents occurring earlier in PCs than in MCs-was absent among males. Conciliatory tendency is higher for Yakushima macaque males (0.31) versus that in studies on the other subspecies Macaca fuscata. We discuss differences in the behavioral ecology of the 2 subspecies, the ecological and social factors that may favor the occurrence of reconciliation, and the possible benefits that males gain from grooming exchange and reconciliation.
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