The organization of collective group movements in wild Barbary macaques (Macaca sylvanus): social structure drives processes of group coordination in macaques

Social animals have to coordinate activities and collective movements to benefit from the advantages of group living. Animals in large groups maintain cohesion by self-organization processes whereas in smaller groups consensus decisions can be reached. Where consensus decisions are relevant leadership may emerge. Variation in the organization of collective movements has been linked to variation in female social tolerance among macaque species ranging from despotic to egalitarian. Here we investigated the processes underlying group movements in a wild macaque species characterized by a degree of social tolerance intermediate to previously studied congeneric species. We focused on processes before, during and after the departure of the first individual. To this end, we observed one group of wild Barbary macaques (Macaca sylvanus) in the Middle Atlas, Morocco using all-occurrence behaviour sampling of 199 collective movements. We found that initiators of a collective movement usually chose the direction in which more individuals displayed pre-departure behavior. Dominant individuals contributed to group movements more than subordinates, especially juveniles, measured as frequencies of successful initiations and pre-departure behaviour. Joining was determined by affiliative relationships and the number of individuals that already joined the movement (mimetism). Thus, in our study group partially shared consensus decisions mediated by selective mimetism seemed to be prevalent, overall supporting the suggestion that a species’ social style affects the organization of group movements. As only the most tolerant species show equally shared consensus decisions whereas in others the decision is partially shared with a bias to dominant individuals the type of consensus decisions seems to follow a stepwise relation. Joining order may also follow a stepwise, however opposite, relationship, because dominance only determined joining in highly despotic, but not in intermediate and tolerant species

Social Bonds Enhance Reproductive Success in Male Macaques

SummaryFor animals living in mixed-sex social groups, females who form strong social bonds with other females live longer and have higher offspring survival [1–3]. These bonds are highly nepotistic, but sometimes strong bonds may also occur between unrelated females if kin are rare [2, 3] and even among postdispersal unrelated females in chimpanzees and horses [4, 5]. Because of fundamental differences between the resources that limit reproductive success in females (food and safety) and males (fertilizations), it has been predicted that bonding among males should be rare and found only for kin and among philopatric males [6] like chimpanzees [7–9]. We studied social bonds among dispersing male Assamese macaques (Macaca assamensis) to see whether males in multimale groups form differentiated social bonds and whether and how males derive fitness benefits from close bonds. We found that strong bonds were linked to coalition formation, which in turn predicted future social dominance, which influenced paternity success. The strength of males' social bonds was directly linked to the number of offspring they sired. Our results show that differentiated social relationships exert an important influence on the breeding success of both sexes that transcends contrasts in relatedness

Social Relationships in Free-Ranging Male Macaca arctoides

Macaque social relationships differ greatly between species. Based on captive studies that focus mainly on females, researchers have classified stumptail macaque (Macaca arctoides) social relationships as tolerant, as indicated by a high rate of affiliation, frequent aggression, and symmetrical conflicts. To accumulate more data on male social relationships, which are relatively understudied, and to generate comparative data, we investigated male social relationships in a provisioned group of 68 free-ranging, naturally dispersing stumptail macaques in southern Thailand. We collected continuous focal animal and ad libitum data on 7 adult and 2 subadult males, recording social behavior during 283 contact hours between December 2006 and March 2007. Stumptail macaques of this population were less tolerant than predicted based on previous studies on captive groups: Rates of spatial proximity, affiliation, and aggression were low, most males directed affiliative behavior toward higher-ranking males, and conflicts were generally of low intensity and relatively asymmetrical. Thus, male stumptail macaques of the focal group appear to differ in their social style from a previous study of a captive group that mainly comprised of females. In some traits, they are even more intolerant than rhesus macaques, an intensively studied intolerant macaque species. We also compare our data on stumptail macaque males to those on other male macaques, but available data are too sparse to draw final conclusions

The evolution of cooperative turn-taking in animal conflict

<p>Abstract</p> <p>Background</p> <p>A fundamental assumption in animal socio-ecology is that animals compete over limited resources. This view has been challenged by the finding that individuals might cooperatively partition resources by "taking turns". Turn-taking occurs when two individuals coordinate their agonistic behaviour in a way that leads to an alternating pattern in who obtains a resource without engaging in costly fights. Cooperative turn-taking has been largely ignored in models of animal conflict and socio-ecological models that explain the evolution of social behaviours based only on contest and scramble competition. Currently it is unclear whether turn-taking should be included in socio-ecological models because the evolution of turn-taking is not well understood. In particular, it is unknown whether turn-taking can evolve when fighting costs and assessment of fighting abilities are not fixed but emerge from evolved within-fight behaviour. We address this problem with an evolutionary agent-based model.</p> <p>Results</p> <p>We found that turn-taking evolves for small resource values, alongside a contest strategy that leads to stable dominance relationships. Turn-taking leads to egalitarian societies with unclear dominance relationships and non-linear dominance hierarchies. Evolutionary stability of turn-taking emerged despite strength differences among individuals and the possibility to evolve within-fight behaviour that allows good assessment of fighting abilities. Evolutionary stability emerged from frequency-dependent effects on fitness, which are modulated by feedbacks between the evolution of within-fight behaviour and the evolution of higher-level conflict strategies.</p> <p>Conclusions</p> <p>Our results reveal the impact of feedbacks between the evolution of within-fight behaviour and the evolution of higher-level conflict strategies, such as turn-taking. Similar feedbacks might be important for the evolution of other conflict strategies such as winner-loser effects or coalitions. However, we are not aware of any study that investigated such feedbacks. Furthermore, our model suggests that turn-taking could be used by animals to partition low value resources, but to our knowledge this has never been tested. The existence of turn-taking might have been overlooked because it leads to societies with similar characteristics that have been expected to emerge from scramble competition. Analyses of temporal interaction patterns could be used to test whether turn-taking occurs in animals.</p

Recruitment and monitoring behaviors by leaders predict following in wild Barbary macaques (Macaca sylvanus)

For group-living animals it is essential to maintain the cohesiveness of the group when traveling. Individuals have to make an accurate decision about where and when to move. Communication before and during the departure of the first individual may play a crucial role in synchronizing a collective movement. We hypothesized that individuals in a wild primate group use signals or cues prior to and after departure to achieve collective movements. With two observers we used all-occurrences behavior sampling of collective movements in a group of wild Barbary macaques (Macaca sylvanus) in the Middle Atlas, Morocco. The number of individuals displaying pre-departure behavior predicted the success of an initiation of a collective movement. Pauses of the first departing individual after departure enhanced following behavior and might have served as recruitment signal. However, the opposite was the case for back-glancing, which functions as a monitoring signal in other species. Because in our study frequently back-glancing individuals were also less socially integrated, back glances may better be interpreted as indicators of hesitation and insecurity. To successfully initiate a collective movement, it seemed to be sufficient for a socially integrated group member to take action when other group members signal their willingness prior to departure and to occasionally wait for the group while movin

The evolution of social philopatry in female primates

The transition from solitary life to sociality is considered one of the major transitions in evolution. In primates, this transition is currently not well understood. Traditional verbal models appear insufficient to unravel the complex interplay of environmental and demographic factors involved in the evolution of primate sociality, and recent phylogenetic reconstructions have produced conflicting results. We therefore analyze a theoretical model for the evolution of female social philopatry that sheds new light on the question why most primates live in groups. In individual-based simulations, we study the evolution of dispersal strategies of both resident females and their offspring. The model reveals that social philopatry can evolve through kin selection, even if retention of offspring is costly in terms of within-group resource competition and provides no direct benefits. Our model supports the role of predator avoidance as a selective pressure for group-living in primates, but it also suggests that a second benefit of group-living, communal resource defense, might be required to trigger the evolution of sizable groups. Lastly, our model reveals that seemingly small differences in demographic parameters can have profound effects on primate social evolution

Quantifying within-group variation in sociality—covariation among metrics and patterns across primate groups and species

It has long been recognized that the patterning of social interactions within a group can give rise to a social structure that holds very different places for different individuals. Such within-group variation in sociality correlates with fitness proxies in fish, birds, and mammals. Broader integration of this research has been hampered by the lack of agreement on how to integrate information from a plethora of dyadic interactions into individual-level metrics. As a step towards standardization, we collected comparative data on affinitive and affiliative interactions from multiple groups each of five species of primates to assess whether the same aspects of sociality are measured by different metrics and indices. We calculated 16 different sociality metrics used in previous research and thought to represent three different sociality concepts. We assessed covariation of metrics within groups and then summarized covariation patterns across all 15 study groups, which varied in size from 5 to 41 adults. With some methodological and conceptual caveats, we found that the number of weak ties individuals formed within their groups represented a dimension of sociality that was largely independent from the overall number of ties as well as from the number and strength of the strong ties they formed. Metrics quantifying indirect connectedness exhibited strong covariation with strong tie metrics and thus failed to capture a third aspect of sociality. Future research linking affiliation and affinity to fitness or other individual level outcomes should quantify inter-individual variation in three aspects: the overall number of ties, the number of weak ties, and the number or strength of strong ties individuals form, after taking into account effects of social network density. Significance statement: In recent years, long-term studies of individually known animals have revealed strong correlations between individual social bonds and social integration, on the one hand, and reproductive success and survival on the other hand, suggesting strong natural selection on affiliative and affinitive behavior within groups. It proved difficult to generalize from these studies because they all measured sociality in slightly different ways. Analyzing covariation between 16 previously used metrics identified only three rather independent dimensions of variation. Thus, different studies have tapped into the same biological phenomenon. How individuals are weakly connected within their group needs further attention.Peer Reviewe

The evolution of between-sex bonds in primates

Social bonds can be a way for individuals to gain access to crucial resources and services that cannot be taken by force and are therefore subject to leverage. Bonds between the sexes can provide access to services that are specific to the other sex. Females exert leverage over males in terms of mating access, males have leverage over females in terms of the service protection, and both sexes exert leverage over the other sex in terms of tolerance and agonistic support. While mating access can be coerced in some circumstances, most services cannot be forced. Here, we use theoretical considerations to explore when sources of leverage over the opposite sex lead to between-sex bonds. Focussing on primates living in multi-male multi-female groups, we predict that leverage over the other sex will be higher, when 1) the receiver benefits on average more than the provider, 2) receivers cannot share the resource, and 3) the resource is rare and valuable. If these conditions are fulfilled, and given the mutual natur