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In long-lived organisms, experience can accumulate with age, such that older individuals may act as repositories of ecological and social knowledge. Such knowledge is often beneficial and can spread via social transmission, leading to the expectation that ageing individuals will remain socially well-integrated. However, social ageing involves multiple processes that modulate the relationship between age and social connectivity in complex ways. We developed a generative model to explore how social ageing may drive changes in social network position and shape older individuals' capacity to transmit knowledge to others. We further employ novel hypernetwork analyses that capture higher-order interactions (i.e. involving ≥ 3 participants) to reveal potential relationships between age and sociality that conventional dyadic networks may overlook. We find that older individuals in our simulations effectively facilitate transmission across a range of scenarios, especially when transmission resembles a complex contagion or when social selectivity (i.e. prioritization of key relationships) rapidly emerges with age. These patterns result from the formation of tight-knit sets of older associates that co-occur in multiple groups, thereby reinforcing one another's capacity to transmit knowledge. Our findings suggest key avenues for future empirical work and illustrate the use of hypernetworks in advancing the study of social behaviour. This article is part of the discussion meeting issue 'Understanding age and society using natural populations'. [ABSTRACT FROM AUTHOR]

Social relationships among animals emerge from interactions in multiple ecological and social situations. However, we seldom ask how each situation contributes to the global structure of a population, and whether different situations contribute different information about social relationships and the position of individuals within the social fabric. Griffon vultures (Gyps fulvus) interact socially in multiple situations, including communal roosting, joint flights, and co‐feeding. These social interactions can influence population‐level outcomes, such as disease transmission and information sharing that determine survival and response to changes. We examined the unique contribution of each social and ecological situation to the social structure of the population and individuals' positions within the overall social network using high‐resolution GPS tracking. We found that the number of individuals each vulture interacted with (degree) was best predicted by diurnal interactions—both during flights and on the ground (such as when feeding). However, the strength of social bonds, that is, the number of interactions an individual had (strength), was best predicted by interactions on the ground—both during the day (e.g., while feeding) and at night (e.g., while roosting) but not by interactions while flying. Thus, social situations differ in their impact on the relationships that individuals form. By incorporating the ecological situations in which social interactions occur we gain a more complete view of how social relationships are formed and which situations are important for different types of interactions. [ABSTRACT FROM AUTHOR]

Individual variation in complex social behavioral traits, like primate grooming, can be influenced by the characteristics of the individual and those of its social group. To better grasp this complexity, social network analysis can be used to quantify direct and indirect grooming relationships. However, multi-group social network studies remain rare, despite their importance to disentangle individual from group-level trait effects on grooming strategies. We applied social network analysis to grooming data of 22 groups of zoo-housed bonobos and investigated the impact of three individual (sex, age, and rearing-history) and two group-level traits (group size and sex ratio) on five social network measures (out-strength, in-strength, disparity, affinity, and eigenvector centrality). Our results showed age-effects on all investigated measures: for females, all measures except for affinity showed quadratic relationships with age, while in males, the effects of age were more variable depending on the network measure. Bonobos with atypical rearing histories showed lower out-strength and eigenvector centrality, while in-strength was only impacted by rearing history in males. Group size showed a negative association with disparity and eigenvector centrality, while sex ratio did not influence any of the investigated measures. Standardization for group size did not impact the effects of sex and age, indicating the robustness of these findings. Our study provides comprehensive insights into the complexity of grooming behavior in zoo-housed bonobos, and underlines the importance of multi-group analyses for the generalizability of social network analysis results for species as a whole. [ABSTRACT FROM AUTHOR]

Aging brings about notable changes in sociality, with an increasing focus on essential partners in both humans and nonhuman primates. Several studies have shown that older nonhuman primates have fewer social partners and shift their types of interactions. The majority of these studies, however, involved only female individuals. Much less is known about the trajectory of social aging in males. We collected 2180 h of focal observation data in a large age‐heterogeneous sample of 34 male and 50 female Barbary macaques (Macaca sylvanus; age range 5–30 years) living in two social groups at the outdoor enclosure at La Forêt des Singes in Rocamadour (France). To track age‐related changes in social engagement of both sexes, we used classical behavioral and social network analysis to measure age‐related variation in the number of partners, the frequency of affiliative interactions, and the position in the social network (eigenvector centrality and local clustering coefficient). We found that females were more central in most social network metrics than males, that is, had more social partners and were more engaged in affiliative interactions than males. We did not find notable sex differences with age: both sexes showed a similar decline in social activity and energy‐demanding activities like jumping or running. Our results thereby extend our knowledge of older nonhuman primates' social life and emphasize that age can have a similar impact on female and male social behavior. Research Highlights: We used social network analysis, as well as analyses of gregariousness, activity patterns and social interest, to test the hypothesis that male Barbary macaques show more variable social aging trajectories compared to females.The analysis of 2180 h of focal observations and scan data from 84 subjects aged 5‐30 years showed that in both sexes, old subjects were less sociable and less physically active, while social interest remained constant.Male aging patterns in this population did not differ substantially from that of females, with a slightly earlier onset of peak social activity in females than in males who hold the most central position when in their prime. [ABSTRACT FROM AUTHOR]

The formalization of multilayer networks allows for new ways to measure sociality in complex social systems, including groups of animals. The same mathematical representation and methods are widely applicable across fields and study systems, and a network can represent drastically different types of data. As such, in order to apply analyses and interpret the results in a meaningful way the researcher must have a deep understanding of what their network is representing and what parts of it are being measured by a given analysis. Multilayer social networks can represent social structure with more detail than is often present in single layer networks, including multiple "types" of individuals, interactions, or relationships, and the extent to which these types are interdependent. Multilayer networks can also encompass a wider range of social scales, which can help overcome complications that are inherent to measuring sociality. In this paper, I dissect multilayer networks into the parts that correspond to different components of social structures. I then discuss common pitfalls to avoid across different stages of multilayer network analyses—some novel and some that always exist in social network analysis but are magnified in multi-layer representations. This paper serves as a primer for building a customized toolkit of multilayer network analyses, to probe components of social structure in animal social systems. [ABSTRACT FROM AUTHOR]

Scale remains a foundational concept in ecology. Spatial scale, for instance, has become a central consideration in the way we understand landscape ecology and animal space use. Meanwhile, scale-dependent social processes can range from fine-scale interactions to co-occurrence and overlapping home ranges. Furthermore, sociality can vary within and across seasons. Multilayer networks promise the explicit integration of the social, spatial, and temporal contexts. Given the complex interplay of sociality and animal space use in heterogeneous landscapes, there remains an important gap in our understanding of the influence of scale on animal social networks. Using an empirical case study, we discuss ways of considering social, spatial, and temporal scale in the context of multilayer caribou social networks. Effective integration of social and spatial processes, including biologically meaningful scales, within the context of animal social networks is an emerging area of research. We incorporate perspectives that link the social environment to spatial processes across scales in a multilayer context. [ABSTRACT FROM AUTHOR]

We provide an overview of the most commonly used social network measures in animal research for static networks or time‐aggregated networks.For each of these measures, we provide clear explanations as to what they measure, we describe their respective variants, we underline the necessity to consider these variants according to the research question addressed, and we indicate considerations that have not been taken so far.We provide a guideline indicating how to use them depending on the data collection protocol, the social system studied and the research question addressed. Finally, we inform about the existent gaps and remaining challenges in the use of several variants and provide future research directions. [ABSTRACT FROM AUTHOR]

Because of the nature of social interaction or association data, when testing hypotheses using social network data it is common for network studies to rely on permutations to control for confounding variables, and to not also control for them in the fitted statistical model. This can be a problem because it does not adjust for any bias in effect sizes generated by these confounding effects, and thus the effect sizes are not informative in the presence of confounding variables.We implemented two network simulation examples and analysed an empirical dataset to demonstrate how relying solely on permutations to control for confounding variables can result in highly biased effect size estimates of animal social preferences that are uninformative when quantifying differences in behaviour.Using these simulations, we show that this can sometimes even lead to effect sizes that have the wrong sign and are thus the effect size is not biologically interpretable. We demonstrate how this problem can be addressed by controlling for confounding variables in the statistical dyadic or nodal model.We recommend this approach should be adopted as standard practice in the statistical analysis of animal social network data. [ABSTRACT FROM AUTHOR]

In social species, network centralities of group members shape social transmission and other social phenomena. Different factors have been found to influence the measurement of social networks, such as data collection and observation methods. In this study, we collected data on adults and juveniles and examined the effect of data collection method (ad libitum sampling vs. focal animal sampling) and observation method (interaction—grooming; play—vs. association—arm‐length; 2 m; 5 m proximities—) on social networks in wild vervet monkeys. First, we showed using a bootstrapping method, that uncertainty of ad libitum grooming and play matrices were lesser than uncertainty of focal matrices. Nevertheless, grooming and play networks constructed from ad libitum and focal animal sampling were very similar and highly correlated. We improved the certainty of both grooming and play networks by pooling focal and ad libitum matrices. Second, we reported a high correlation between the proximity arm‐length network and the focal grooming one making an arm‐length proximity network a reasonable proxy for a grooming one in vervet monkeys. However, we did not find such a correlation between proximity networks and the play one. Studying the effects of methodological issues as data collection and observation methods can help improve understanding of what shapes social networks and which data collection method to choose to study sociality. HIGHLIGHTS: We compared social networks constructed through different data collection and observation methods in a wild primate species.We found that ad libitum networks were less uncertain than focal networks.Arm‐length proximity network can be used as a proxy for a grooming network but not for a play one. [ABSTRACT FROM AUTHOR]

The requirements of living in social groups, and forming and maintaining social relationships are hypothesized to be one of the major drivers behind the evolution of cognitive abilities. Most empirical studies investigating the relationships between sociality and cognition compare cognitive performance between species living in systems that differ in social complexity. In this review, we ask whether and how individuals benefit from cognitive skills in their social interactions. Cognitive abilities, such as perception, attention, learning, memory, and inhibitory control, aid in forming and maintaining social relationships. We investigate whether there is evidence that individual variation in these abilities influences individual variation in social relationships. We then consider the evolutionary consequences of the interaction between sociality and cognitive ability to address whether bi-directional relationships exist between the two, such that cognition can both shape and be shaped by social interactions and the social environment. In doing so, we suggest that social network analysis is emerging as a powerful tool that can be used to test for directional causal relationships between sociality and cognition. Overall, our review highlights the importance of investigating individual variation in cognition to understand how it shapes the patterns of social relationships. [ABSTRACT FROM AUTHOR]

The social relationships that individuals experience at different life stages have a non‐negligible influence on their lives, and this is particularly true for group living animals. The long lifespan of many primates makes it likely that these animals have various tactics of social interaction to adapt to complex changes in environmental or physical conditions. The different strategies used in social interaction by individuals at different life stages, and whether the position (central or peripheral) or role (initiator or recipient) of an individual in the group social network changes with age, are intriguing questions that remain to be investigated. We used social network analysis to examine age‐related differences in social interaction patterns, social roles, and social positions in three affiliative social networks (approach, allogrooming, and social play) in a group of wild rhesus macaques (Macaca mulatta). Our results showed that social interaction patterns of rhesus macaques differ between age classes in the following ways: i) young individuals tend to allocate social time to a high number of groupmates, older individuals prefer to focus on fewer, specific partners; ii) as they grow older, individuals tend to be recipients in approach interactions and initiators in grooming interactions; and iii) regardless of the different social interaction strategies, individuals of all ages occupy a central position in the group. These results reveal a possible key role played by immature individuals in group social communication, a little‐explored issue which deserves closer investigation in future research. [ABSTRACT FROM AUTHOR]

Simple Summary: The maintenance and stability of social experience is an especially important element of the captive welfare of zoo-housed species. The population of critically endangered Livingstone's fruit bats (Pteropus livingstonii) resident at Jersey Zoo display a complex social structure of affiliative and aggressive interactions. Subgroups defined by individual characteristics contribute in different ways to this structure. Social information, illuminated through the use of social network analysis techniques, could be used in the future to promote social stability and safeguard individual welfare when making evidence-based husbandry decisions. Social network analysis has been highlighted as a powerful tool to enhance the evidence-based management of captive-housed species through its ability to quantify the social experience of individuals. We apply this technique to explore the social structure and social roles of 50 Livingstone's fruit bats (Pteropus livingstonii) housed at Jersey Zoo, Channel Islands, through the observation of associative, affiliative, and aggressive interactions over two data collection periods. We implement binomial mixture modelling and characteristic-based assortment quantification to describe the complexity and organisation of social networks, as well as a multiple regression quadratic assignment procedural (MRQAP) test to analyse the relationship between network types. We examine the effects of individual characteristics (i.e., sex, age, and dominance rank) on social role by fitting models to explain the magnitude of node metrics. Additionally, we utilize a quadratic assignment procedural (QAP) test to assess the temporal stability of social roles over two seasons. Our results indicate that P. livingstonii display a non-random network structure. Observed social networks are positively assorted by age, as well as dominance rank. The frequency of association between individuals correlates with a higher frequency of behavioural interactions, both affiliative and aggressive. Individual social roles remain consistent over ten months. We recommend that, to improve welfare and captive breeding success, relationships between individuals of similar ages and dominance levels should be allowed to persist in this group where possible, and separating individuals that interact frequently in an affiliative context should be avoided. [ABSTRACT FROM AUTHOR]