Group-level cooperation often poses a social dilemma in which joint action may be difficult to achieve. Theoretical models and experimental work on humans show that social incentives, such as punishment of defectors and rewarding of cooperators, can promote cooperation in groups of unrelated individuals. Here, we demonstrate that these processes can operate in a non-human animal species, and be used to effectively promote the production of a public good. We took advantage of the fact that intergroup fights in vervet monkeys ( Chlorocebus aethiops pygerythrus ) are characterized by episodes of intergroup aggression with pauses in-between. During pauses, females selectively groomed males that had participated in the previous aggressive episode, but aggressed male group members that had not. In subsequent (i.e. future) episodes, males who had received either aggression or grooming participated above their personal base-line level. Therefore, female–male aggression and grooming both appear to function as social incentives that effectively promote male participation in intergroup fights. Importantly, females stood to gain much from recruiting males as the probability of winning intergroup fights was dependent on the number of active participants, relative to the number of fighters in the opposing group. Furthermore, females appear to maximize the benefits gained from recruiting males as they primarily used social incentives where and when high-quality food resources, which are the resources primarily limiting to female fitness, were at stake.

In social species, fighting in intergroup conflicts is one of the riskiest cooperative activities group members engage in, particularly for individuals of the smaller sex. In a number of species, female group members are significantly smaller than males and as a result, the costs associated with intergroup aggression outweigh the potential benefits, and females avoid participating. Studies conducted on species in which females are active participants have consistently found that they fight to defend access to food resources and that high-ranking females tend to be more active than low-rankers. However, additional factors may modulate the costs and benefits of participation, creating differences among individuals and variability within individuals over time. In this study, we investigate a number of costs and benefits that potentially affect female vervet monkey (Chlorocebus aethiops pygerythrus) participation in intergroup conflicts. We observed the participation of 35 females in 3 groups, during 115 intergroup conflicts. Our findings suggest that female vervet monkeys defend access to valuable food resources, as well as to areas that are intensely used in the long-term; however, rank also influenced female participation indicating that the potential benefits gained from resource defence vary with one's priority of access to these resources. We find that females were more likely to participate aggressively when they did not have an infant, and when they received more male support throughout the intergroup conflict, suggesting these factors influence the perceived risk, or costs, of intergroup aggression. Because we observed considerable temporal variability in both the proportion of female group members with infants, and the number and identity of male group members (i.e. amount of male support provided), the relative fighting ability of neighbouring groups will inevitably fluctuate. Thus, our findings may help explain the lack of stable intergroup dominance relationships observed in many studies of intergroup conflict.

In numerous social species, males direct aggression towards female group members during intergroup fights, and this behaviour is commonly thought to function as mate guarding, even though males often target non-receptive females. In studying intergroup fights in a wild population of vervet monkeys, we found that male intragroup aggression was primarily directed towards individuals who had either just finished exhibiting, or were currently attempting to instigate intergroup aggression. Targeted females were less likely to instigate intergroup aggression in the future, indicating that male intragroup aggression functioned as coercion (when directed towards those who were currently trying to instigate a fight) and punishment (when directed towards those who had recently fought). These manipulative tactics effectively prevented intergroup encounters from escalating into fights and often de-escalated ongoing conflicts. Males who were likely sires were those most likely to use punishment/coercion, particularly when they were wounded, and, therefore, less able to protect vulnerable offspring should a risky intergroup fight erupt. This work, along with our previous finding that females use punishment and rewards to recruit males into participating in intergroup fights, highlights the inherent conflict of interest that exists between the sexes, as well as the role that social incentives can play in resolving this conflict. Furthermore, unlike other studies which have found punishment to be used asymmetrically between partners, these works represent a novel example of reciprocal punishment in a non-human animal.

Males in a number of group-living species fight in intergroup conflicts to defend access to food resources, a seemingly paradoxical behaviour, given that this resource does not usually limit male fitness directly. We investigated the mechanism(s) driving apparent male food defence in wild vervet monkeys (Chlorocebus aethiops pygerythrus) by testing the effect that female resource access, and female audience size and activity had on the response of focal males during simulated intergroup encounters. Males do not appear to defend food to increase the reproductive success of female group members because their response was not influenced by the presence of provisioning boxes that only females could access. Female audience size was also unimportant, suggesting males do not participate in intergroup encounters to advertise their quality to potential mates. However, focal males almost always followed/supported female group members who initiated an approach towards simulated intruders, supporting that male participation largely functions to gain status as a cooperative group member, and that apparent male food defence in this species arises as a by-product of intersexual cooperation. Our study highlights that considering audience composition and activity can reveal the presence of social incentives and illuminate the evolutionary mechanism(s) promoting joint action in intergroup aggression.

Close proximity and social grooming are important bonding mechanisms in primates. These behaviors show the social structure of a species and many studies have found positive correlations between the degree of kinship and grooming and proximity. We used 1 year of data collected via instantaneous scan sampling on a large "supertroop" of Colobus angolensis ruwenzorii at Lake Nabugabo, Uganda, to examine partner preferences for grooming and nearest neighbors in each age-sex class. Little is known about this species, so we based our hypotheses on congeners. Of the five species of black-and-white colobus, data on sex-biased dispersal patterns are available for three (C. guereza, C. vellerosus, and C. polykomos), all of which show male-biased dispersal with occasional female dispersal. We thus predicted that female C. a. ruwenzorii would be more strongly bonded than males, showing greater proximity and grooming. We did not expect bonding between the sexes since congeners do not show this pattern. We found that among adult dyads, males and females were more likely to be found in loose proximity, and to groom, than would be expected given group composition. Conversely, both males and females had relatively weak same-sex relationships. Between the sexes, adult males had higher proximity and grooming indices with adult females without infants than with females with infants. These observations indicate that this subspecies is cross-bonded and that both sexes may disperse. Furthermore, our findings suggest that the social organization and social structure of C. a. ruwenzorii differ greatly from other black-and-white colobus species.

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<b><i>Introduction:</i></b> Colobine monkeys are specialized folivores that use foregut fermentation to digest leaves. The slow process of fermentation forces them to spend a lot of time resting and to minimize their energy expenditure to subsist on a lower-quality diet. <b><i>Methods:</i></b> We recorded the diet and activity budget of <i>Colobus angolensis ruwenzorii</i>, which form a three-tiered multi-level society, at Lake Nabugabo, Uganda, over 12 months using scan sampling on adults and subadults, to determine whether they utilize the energy minimization strategy typical of colobines. <b><i>Results:</i></b> We found that the annual diet was primarily comprised of high-quality food resources (young leaves 65% and fruit 31%), and fruits were the only plant part the monkeys<i></i>selected when available. Both the fruits and young leaves of some species were preferred food items in some months, and mature leaf consumption correlated negatively with preferred food availability. Mature leaves appear to be a fallback food for this population but are rarely relied upon (3%). The <i>C. a. ruwenzorii</i>at Nabugabo spent less time resting (40%) and more time moving (25%) than is typical for other species of black-and-white colobus. <b><i>Discussion/Conclusion:</i></b> The high-quality diet of this population appears to allow them to utilize an energy maximization strategy. Their reliance on food items that tend to be clumped in space and time likely explains the frequent fission-fusion behaviour that we observe between core units. Our findings demonstrate that the foraging strategies of colobines may be more flexible than was previously thought and illustrate how food availability and distribution can impact primate social organization.

Foraging animals need to quickly assess the costs and benefits of different foraging decisions, including resource quantity, quality, preference, ease of access, dispersion, distance, and predation risk. Social animals also need to take social context into account and adapt foraging strategies that maximize net resource intake and minimize contest competition with conspecifics. We used an experimental approach to investigate how social context impacts wild vervet monkey (Chlorocebus pygerythrus) foraging decisions in a multi-destination pentagon array. We baited four platforms with less-preferred corn and one platform with a larger, preferred resource (half banana) that required handling time. We ran over 1,000 trials and found that when monkeys foraged alone, they usually took the path that minimized travel distance but prioritized the preferred-food platform when in competition. However, the foraging strategy chosen by low-ranking individuals depended on the handling skill of the decision maker (i.e., time it would take them to retrieve the banana), the relative rank of their audience members (i.e., who has priority-of-access to resources), and the distance audience members were from the experiment site (i.e., their travel time). When the risk of being displaced by a dominant competitor was low (because they were far away and/or because the decision-maker was skilled in retrieving the banana), low-ranking individuals chose a route that minimized travel costs. Conversely, when the risk of losing food to a dominant competitor was high, decision-makers rushed for the preferred-food platform at the onset of the trial. When the risk of displacement was moderate because a dominant audience member was at least 50 m away, low-ranking individuals partly prioritized the preferred-food platform but took the time to stop for one platform of corn on the way. This strategy increased the total amount of food obtained during the trial. These findings suggest that lower-ranking individuals, who experienced high contest competition at the foraging experiment, calculated the risk of being displaced by a dominant competitor when making foraging decisions. This experiment demonstrates that vervets go through a complex decision-making process that simultaneously considers the profitability of different foraging decisions and their social context.