Cooperative breeding and the emergence of multilevel societies in birds

Multilevel societies (MLSs), where social levels are hierarchically nested within each other, are considered one of the most complex forms of animal societies. Although thought to mainly occur in mammals, it is suggested that MLSs could be under-detected in birds. Here we propose that the emergence of MLSs could be common in cooperatively breeding birds, as both systems are favoured by similar ecological and social drivers. We first investigate this proposition by systematically comparing evidence for multilevel social structure in cooperative and non-cooperative birds in Australia and New Zealand, global hotspots for cooperative breeding. We then analyse non-breeding social networks of cooperatively breeding superb fairy-wrens (Malurus cyaneus) to reveal their structured multilevel society, with three hierarchical social levels that are stable across years. Our results confirm recent predictions that MLSs are likely to be widespread in birds and suggest that these societies could be particularly common in cooperatively breeding birds.

Group Composition and Social Structure of Red-Shanked Doucs (Pygathrix nemaeus) at Son Tra Nature Reserve, Vietnam

Multilevel societies, consisting of multiple one-male multi-female units, are relatively rare among primates, but are more widespread in the odd-nosed colobines than other taxa. Multilevel societies are found particularly in snub-nosed monkeys (<i>Rhinopithecus</i>)<i></i>and have been debated in studies of proboscis monkeys (<i>Proboscis</i>). While it has been suggested that douc langurs (<i>Pygathrix</i>) may also form multilevel societies, the limited data available make the details of their social organization unclear. We aimed to establish a more comprehensive picture of the social organization of red-shanked doucs (<i>Pygathrix nemaeus</i>) and to address the question of whether this species forms multilevel societies, specifically collections of multiple distinct one-male units hereafter termed “bands.” We collected 259 h of behavioral data at Son Tra Nature Reserve in Vietnam from February 2010 to May 2011. The mean band size was approximately 18 individuals. Bands were comprised of approximately 2.7 units, and each unit contained approximately 6.5 individuals. Units had an average sex ratio of 1.0:1.6. We observed fission and fusion behaviors which were not correlated with phenological or weather measures. Activity budget data showed that fission and fusion behaviors between units were positively correlated with activity. Both vocalizations and vigilance increased when units engaged in fission. Based on this evidence, <i>P. nemaeus</i> at Son Tra Nature Reserve appear to engage in daily fission-fusion activity which does not vary between seasons. Additionally, our data suggest that these primates may form multilevel societies made up of distinct units. However, future data including proximity pattern analyses are necessary for confirmation.

Aerial drone observations identified a multilevel society in feral horses

The study of non-human multilevel societies can give us insights into how group-level relationships function and are maintained in a social system, but their mechanisms are still poorly understood. The aim of this study was to apply spatial association data obtained from drones to verify the presence of a multilevel structure in a feral horse society. We took aerial photos of individuals that appeared in pre-fixed areas and collected positional data. The threshold distance of the association was defined based on the distribution pattern of the inter-individual distance. The association rates of individuals showed bimodality, suggesting the presence of small social organizations or “units”. Inter-unit distances were significantly smaller than those in randomly replaced data, which showed that units associate to form a higher-level social organization or “herd”. Moreover, this herd had a structure where large mixed-sex units were more likely to occupy the center than small mixed-sex units and all-male-units, which were instead on the periphery. These three pieces of evidence regarding the existence of units, unit association, and stable positioning among units strongly indicated a multilevel structure in horse society. The present study contributes to understanding the functions and mechanisms of multilevel societies through comparisons with other social indices and models as well as cross-species comparisons in future studies.

Beyond the group: how food, mates, and group size influence intergroup encounters in wild bonobos

In social-living animals, interactions between groups are frequently agonistic, but they can also be tolerant and even cooperative. Intergroup tolerance and cooperation are regarded as a crucial step in the formation of highly structured multilevel societies. Behavioral ecological theory suggests that intergroup tolerance and cooperation can emerge either when the costs of hostility outweigh the benefits of exclusive resource access or when both groups gain fitness benefits through their interactions. However, the factors promoting intergroup tolerance are still unclear due to the paucity of data on intergroup interactions in tolerant species. Here, we examine how social and ecological factors affect the onset and termination of intercommunity encounters in two neighboring communities of wild bonobos, a species exhibiting flexible patterns of intergroup interactions, at Kokolopori Bonobo Reserve, Democratic Republic of the Congo. We recorded the timing and location of intercommunity encounters and measured fruit abundance and distribution, groups’ social characteristics, and space-use dynamics over a 19-month period. We found that intercommunity tolerance was facilitated by a decrease in feeding competition, with high fruit abundance increasing the likelihood of communities to encounter, and high clumpiness of fruit patches increasing the probability to terminate encounters likely due to increased contest. In addition, the possibility for extra-community mating, as well as the potential benefits of more efficient foraging in less familiar areas, reduced the probability that the communities terminated encounters. By investigating the factors involved in shaping relationships across groups, this study contributes to our understanding of how animal sociality can extend beyond the group level.

Convergence and divergence in gesture repertoires as an adaptive mechanism for social bonding in primates

A key challenge for primates living in large, stable social groups is managing social relationships. Chimpanzee gestures may act as a time-efficient social bonding mechanism, and the presence (homogeneity) and absence (heterogeneity) of overlap in repertoires in particular may play an important role in social bonding. However, how homogeneity and heterogeneity in the gestural repertoire of primates relate to social interaction is poorly understood. We used social network analysis and generalized linear mixed modelling to examine this question in wild chimpanzees. The repertoire size of both homogeneous and heterogeneous visual, tactile and auditory gestures was associated with the duration of time spent in social bonding behaviour, centrality in the social bonding network and demography. The audience size of partners who displayed similar or different characteristics to the signaller (e.g. same or opposite age or sex category) also influenced the use of homogeneous and heterogeneous gestures. Homogeneous and heterogeneous gestures were differentially associated with the presence of emotional reactions in response to the gesture and the presence of a change in the recipient's behaviour. Homogeneity and heterogeneity of gestural communication play a key role in maintaining a differentiated set of strong and weak social relationships in complex, multilevel societies.