The social network structure of a wild meerkat population: 2. Intragroup interactions

Knowledge of the structure of networks of social interactions is important for understanding the evolution of cooperation, transmission of disease, and patterns of social learning, yet little is known of how environmental, ecological, or behavioural factors relate to such structures within groups. We observed grooming, dominance, and foraging competition interactions in eight groups of wild meerkats (Suricata suricatta) and constructed interaction networks for each behaviour. We investigated relationships between networks for different social interactions and explored how group attributes (size and sex ratio), individual attributes (tenure of dominants), and ecological factors (ectoparasite load) are related to variation in network structure. Network structures varied within a group according to interaction type. Further, network structure varied predictably with group attributes, individual attributes, and ecological factors. Networks became less dense as group size increased suggesting that individuals were limited in their number of partners. Groups with more established dominant females were more egalitarian in their grooming and foraging competition interactions, but more despotic in their dominance interactions. The distribution of individuals receiving grooming became more skewed at higher parasite loads, but more equitable at low parasite loads. We conclude that the pattern of interactions between members of meerkat groups is not consistent between groups but instead depends on general attributes of the group, the influence of specific individuals within the group, and ecological factors acting on group members. We suggest that the variation observed in interaction patterns between members of meerkat groups may have fitness consequences both for individual group members and the group itself.     

Leading from the front? Social networks in navigating groups

In many group-living animals, leadership by only a fraction of the group members can be important for group navigation. It has been shown that subgroups of informed individuals can steer the remainder of the group without direct communication, resolving conflicts of interest through individual-to-individual interactions. We present a model for the navigation of collectively moving groups that includes preferential interactions between individuals as a way of imposing social network structures, known to be present in many species. We show that effective leadership can occur when leaders do not occupy frontal spatial positions and when navigation tendency is appropriately balanced with social position. Our model also shows that small minorities can dominate movement decisions if they have navigational knowledge combined with influential social network positions. Our findings highlight the mechanistic importance of social networks for the movement decisions of animal groups. We discuss the implications of our research for interpreting empirical observations.     

Shark personalities? Repeatability of social network traits in a widely distributed predatory fish

Interest in animal personalities has generated a burgeoning literature on repeatability in individual traits such as boldness or exploration through time or across different contexts. Yet, repeatability can be influenced by the interactive social strategies of individuals, for example, consistent inter-individual variation in aggression is well documented. Previous work has largely focused on the social aspects of repeatability in animal behaviour by testing individuals in dyadic pairings. Under natural conditions, individuals interact in a heterogeneous polyadic network. However, the extent to which there is repeatability of social traits at this higher order network level remains unknown. Here, we provide the first empirical evidence of consistent and repeatable animal social networks. Using a model species of shark, a taxonomic group in which repeatability in behaviour has yet to be described, we repeatedly quantified the social networks of ten independent shark groups across different habitats, testing repeatability in individual network position under changing environments. To understand better the mechanisms behind repeatable social behaviour, we also explored the coupling between individual preferences for specific group sizes and social network position. We quantify repeatability in sharks by demonstrating that despite changes in aggregation measured at the group level, the social network position of individuals is consistent across treatments. Group size preferences were found to influence the social network position of individuals in small groups but less so for larger groups suggesting network structure, and thus, repeatability was driven by social preference over aggregation tendency.     

The social network structure of a wild meerkat population: 1. Inter-group interactions

Groups of individuals frequently interact with each other, but typically analysis of such interactions is restricted to isolated dyads. Social network analysis (SNA) provides a method of analysing polyadic interactions and is used to analyse interactions between individuals. We use a population of 12 groups (ca. 250 animals) of wild meerkats (Suricata suricatta) to test whether SNA can also be used to describe and elucidate patterns of inter-group interactions. Using data collected over 24 months, we constructed two sets of networks, based on direct encounters between groups and instances of roving males visiting other groups. We analysed replicated networks of each type of interaction to investigate similarities between networks of different social interactions as well as testing their stability over time. The two network types were similar to each other when derived from long-term data, but showed significant differences in structure over shorter timescales where they varied according to seasonal and ecological conditions. Networks for both types of inter-group interaction constructed from data collected over 3 months reliably described long-term (12- and 24-month) patterns of interactions between groups, indicating a stable social structure despite variation in group sizes and sex ratios over time. The centrality of each meerkat group in roving interactions networks was unaffected by the sex ratio of its members, indicating that male meerkats preferentially visit geographically close groups rather than those containing most females. Indeed, the strongest predictors of network structure were spatial factors, suggesting that, in contrast to analyses of intra-group interactions, analyses of inter-group interactions using SNA must take spatial factors into account.     

Network position: a key component in the characterization of social personality types

In recent years, animal social interactions have received much attention in terms of personality research (e.g. aggressive or cooperative interactions). However, other components of social behaviour such as those describing the intensity, frequency, directedness and individual repeatability of interactions in animal groups have largely been neglected. Network analysis offers a valuable opportunity to characterize individual consistency of traits in labile social groups and therein provide novel insights to personality research in ways previously not possible using traditional techniques. Should individual network positions be consistently different between individuals under changing conditions, they might reflect expressions of an individual's personality. Here, we discuss a conceptual framework for using network analyses to infer the presence of individual differences and present a statistical test based on randomization techniques for testing the consistency of network positions in individuals. The statistical tools presented are useful because if particular individuals consistently occupy key positions in social networks, then this is also likely to have consequences for their fitness as well as for that of others in the population. These consequences may be particularly significant since individual network position has been shown to be important for the transmission of diseases, socially learnt information and genetic material between individuals and populations.     

Social attributes and associated performance measures in marmots: bigger male bullies and weakly affiliating females have higher annual reproductive success

Studying the structure of social interactions is fundamental in behavioral ecology as social behavior often influences fitness and thus natural selection. However, social structure is often complex, and determining the most appropriate measures of variation in social behavior among individuals can be difficult. Social network analysis generates numerous, but often correlated, measures of individual connectedness derived from a network of interactions. We used measures of individual connectedness in networks of affiliative and agonistic interactions in yellow-bellied marmots, Marmota flaviventris, to first determine how variance was structured among network measures. Principal component analysis reduced our set of network measures to four “social attributes” (unweighted connectedness, affiliation strength, victimization, and bullying), which revealed differences between patterns of affiliative and agonistic interactions. We then used these extracted social attributes to examine the relationship between an individual’s social attributes and several performance measures: annual reproductive success, parasite infection, and basal stress. In male marmots, bullying was positively associated with annual reproductive success, while in females, affiliation strength was negatively associated with annual reproductive success. No other social attributes were significantly associated with any performance measures. Our study highlights the utility of considering multiple dimensions when measuring the structure and functional consequences of social behavior.     

Loud calls as a mechanism of social coordination in a fission–fusion taxon, the white-bellied spider monkey (Ateles belzebuth)

Spider monkeys (Ateles spp.) live in social groups that exhibit high levels of fission–fusion dynamics, in which group members form subgroups of varying sizes and compositions. Within these fluid societies, how individuals establish contact with dispersed group members with whom they might choose to associate remains unclear. Long-range vocalizations might facilitate interactions between group members and provide a means of social coordination in fission–fusion societies. We evaluated this possibility for one spider monkey vocalization, the loud call, by examining calling behavior, the relationship between loud calls and changes in subgroup size, and the response of individuals to distant calls and playback experiments in a single study group. We found that 82 % of loud calls were emitted within 30 min of a call from a different location, suggesting that individuals frequently emit loud calls in response to the calls of distant group members. Subgroups that emitted loud calls, especially those that responded to distant calls, were much more likely to experience an increase in subgroup size within an hour after calling than those that did not. Animals also approached distant loud calls more than they avoided or ignored these calls. Finally, playbacks of male calls demonstrated that females respond preferentially to the calls of some individuals over others. Taken together, these results provide support for the hypothesis that spider monkey loud calls function to facilitate and initiate interactions between dispersed group members and suggest that vocal signals can play an important role in influencing social interactions in fission–fusion societies.     

Behavioral correlates of non-random mortality among free-ranging female vervet monkeys

Summary Over a 3.5 year period, illness and predation operated in a non-random manner on free-ranging vervet monkeys in Amboseli National Park. As a result, there was no correlation among adult females between dominance rank and reproductive success. Deaths due to illness were concentrated among low-ranking individuals, and appeared to occur as a result of restricted access to food and water during the dry season. In contrast, deaths due to predation were concentrated among high-ranking individuals. The precise cause of such increased vulnerability could not be determined.High-ranking females alarm-called at higher frequencies than low-ranking females, and were also more aggressive than low-ranking females during intergroup encounters. In contrast, low-ranking females were more likely to initiate friendly interactions with the members of other groups. The non-random distribution of causes of mortality suggests that individuals living in the same social group may confront different selective pressures. Perhaps as a result, individuals appear to respond differently to similar social and environmental variables.     

Social network theory: new insights and issues for behavioral ecologists

Until recently, few studies have used social network theory (SNT) and metrics to examine how social network structure (SNS) might influence social behavior and social dynamics in non-human animals. Here, we present an overview of why and how the social network approach might be useful for behavioral ecology. We first note four important aspects of SNS that are commonly observed, but relatively rarely quantified: (1) that within a social group, differences among individuals in their social experiences and connections affect individual and group outcomes; (2) that indirect connections can be important (e.g., partners of your partners matter); (3) that individuals differ in their importance in the social network (some can be considered keystone individuals); and (4) that social network traits often carry over across contexts (e.g., SN position in male–male competition can influence later male mating success). We then discuss how these four points, and the social network approach in general, can yield new insights and questions for a broad range of issues in behavioral ecology including: mate choice, alternative mating tactics, male–male competition, cooperation, reciprocal altruism, eavesdropping, kin selection, dominance hierarchies, social learning, information flow, social foraging, and cooperative antipredator behavior. Finally, we suggest future directions including: (1) integrating behavioral syndromes and SNT; (2) comparing space use and SNS; (3) adaptive partner choice and SNS; (4) the dynamics and stability (or instability) of social networks, and (5) group selection shaping SNS. This contribution is part of the special issue “Social networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau and R. James).     

Social organisation of thornbill-dominated mixed-species flocks using social network analysis

Mixed-species associations are a widespread phenomenon, comprising interacting heterospecific individuals which gain predator, foraging or social benefits. Avian flocks have traditionally been classified as monolithic species units, with species-wide functional roles, such as nuclear, active, passive, or follower. It has also been suggested that flocks are mutualistic interactions, where niches of participating species converge. However the species-level perspective has limited previous studies, because both interactions and benefits occur at the level of the individual. Social network analysis provides a set of tools for quantitative assessment of individual participation. We used mark-resighting methods to develop networks of nodes (colour-marked individuals) and edges (their interactions within flocks). We found that variation in flock participation across individuals within species, especially in the buff-rumped thornbill, encompassed virtually the entire range of variation across all individuals in the entire set of species. For example, female, but not male, buff-rumped thornbills had high network betweenness, indicating that they interact with multiple flocks, likely as part of a female-specific dispersal strategy. Finally, we provide new evidence that mixed-species flocking is mutualistic, by quantifying an active shift in individual foraging niches towards those of their individual associates, with implications for trade-off between costs and benefits to individuals derived from participating in mixed-species flocks. This study is, to our knowledge, the first instance of a heterospecific social network built on pairwise interactions.     

Network structure and parasite transmission in a group living lizard,the gidgee skink, <Emphasis Type="Italic">Egernia stokesii</Emphasis>

Gidgee skinks (Egernia stokesii) form large social aggregations in rocky outcrops across the Flinders Ranges in South Australia. Group members share refuges (rock crevices), which may promote parasite transmission. We measured connectivity of individuals in networks constructed from patterns of common crevice use and observed patterns of parasitism by three blood parasites (Hemolivia, Schellackia and Plasmodium) and an ectoparasitic tick (Amblyomma vikirri). Data came from a 1-year mark-recapture study of four populations. Transmission networks were constructed to represent possible transmission pathways among lizards. Two lizards that used the same refuge within an estimated transmission period were considered connected in the transmission network. An edge was placed between them, directed towards the individual that occupied the crevice last. Social networks, a sub-set of same-day only associations, were small and highly fragmented compared with transmission networks, suggesting that non-synchronous crevice use leads to more transmission opportunities than direct social association. In transmission networks, lizards infested by ticks were connected to more other tick-infested lizards than uninfected lizards. Lizards infected by ticks and carrying multiple blood parasite infections were in more connected positions in the network than lizards without ticks or with one or no blood parasites. Our findings suggest higher levels of network connectivity may increase the risk of becoming infected or that parasites influence lizard behaviour and consequently their position in the network. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau and R. James).     

Reputation is valuable within and outside one’s own social group

To find conditions under which humans cooperate within groups of unrelated individuals has been of major interest in the behavioral sciences. The experimental paradigm for studying potential cooperation in social dilemmas is the public goods game. Here humans regularly fail to sustain a public resource cooperatively. However, the need to maintain good reputation for other social interactions, such as indirect reciprocity, has been identified as an effective mechanism to sustain cooperation in public goods situations. As a side effect of building a good reputation through cooperative actions, an individual provides direct benefits to members of his/her own social group. These benefits could be an incentive to reward a good reputation of group members. Here we show experimentally that building a good reputation through cooperative behavior in a public goods situation is rewarded in future social interactions, not only within ones own social group but also, at a similar level, in other social groups: humans regard cooperative behavior of others as an honest signal irrespective of past direct personal benefits. Reputation gained within as well as outside ones own social group can be a driving force for selfish individuals to cooperate in public goods situations, and thereby sustain any public resource.Communicated by T. Czeschlik     

The bottlenose dolphin community of Doubtful Sound features a large proportion of long-lasting associations

More than 12 studies of different bottlenose dolphin populations, spanning from tropical to cold temperate waters, have shown that the species typically lives in societies in which relationships among individuals are predominantly fluid. In all cases dolphins lived in small groups characterised by fluid and dynamic interactions and some degree of dispersal from the natal group by both sexes. We describe a small, closed population of bottlenose dolphins living at the southern extreme of the species' range. Individuals live in large, mixed-sex groups in which no permanent emigration/immigration has been observed over the past 7 years. All members within the community are relatively closely associated (average half-weight index>0.4). Both male–male and female–female networks of preferred associates are present, as are long-lasting associations across sexes. The community structure is temporally stable, compared to other bottlenose dolphin populations, and constant companionship seems to be prevalent in the temporal association pattern. Such high degrees of stability are unprecedented in studies of bottlenose dolphins and may be related to the ecological constraints of Doubtful Sound. Fjords are low-productivity systems in which survival may easily require a greater level of co-operation, and hence group stability. These conditions are also present in other cetacean populations forming stable groups. We therefore hypothesise that ecological constraints are important factors shaping social interactions within cetacean societies.Communicated by D. Watts     

A test of multiple hypotheses for the function of call sharing in female budgerigars, Melopsittacus undulatus

In many social species group, members share acoustically similar calls. Functional hypotheses have been proposed for call sharing, but previous studies have been limited by an inability to distinguish among these hypotheses. We examined the function of vocal sharing in female budgerigars with a two-part experimental design that allowed us to distinguish between two functional hypotheses. The social association hypothesis proposes that shared calls help animals mediate affiliative and aggressive interactions, while the password hypothesis proposes that shared calls allow animals to distinguish group identity and exclude nonmembers. We also tested the labeling hypothesis, a mechanistic explanation which proposes that shared calls are used to address specific individuals within the sender–receiver relationship. We tested the social association hypothesis by creating four–member flocks of unfamiliar female budgerigars (Melopsittacus undulatus) and then monitoring the birds’ calls, social behaviors, and stress levels via fecal glucocorticoid metabolites. We tested the password hypothesis by moving immigrants into established social groups. To test the labeling hypothesis, we conducted additional recording sessions in which individuals were paired with different group members. The social association hypothesis was supported by the development of multiple shared call types in each cage and a correlation between the number of shared call types and the number of aggressive interactions between pairs of birds. We also found support for calls serving as a labeling mechanism using discriminant function analysis with a permutation procedure. Our results did not support the password hypothesis, as there was no difference in stress or directed behaviors between immigrant and control birds.     

Greater spear-nosed bats give group-distinctive calls

Individually distinctive vocalizations are ubiquitous; however, group distinctive calls have rarely been demonstrated. Under some conditions, selection should favor calls indicating social group membership in animals that forage in groups. Greater spear-nosed bats (Phyllostomus hastatus) give calls that appear to facilitate recognition of social group mates who are unrelated. Females give loud broadband (4–18 kHz) vocalizations termed screech calls when departing on foraging trips and at foraging sites. Screech calls help to establish foraging groups among social group members, and to maintain contact over the long distances they travel while foraging. I test two hypotheses about how screech calls may be structured to convey caller identity. Individual calls may be distinct and group members may learn to recognize each individual's calls and to associate the individual with the social group. Alternatively, groups may give distinct calls and individuals within groups may share call characteristics. To test these hypotheses I conducted multivariate acoustic analysis of multiple calls from 28 bats from three social groups. Although the ubiquity of individually distinctive calls in other taxa makes this result more likely, the results reveal that group calls are highly distinctive. Individual bats within groups are statistically indistinguishable. Calls appear to decrease slightly in frequency as bats age. Call convergence among unrelated group mates implies vocal learning in this species. Received: 28 March 1996 / Accepted after revision: 6 October 1996     

Wildlife contact analysis: emerging methods, questions, and challenges

Recent technological advances, such as proximity loggers, allow researchers to collect complete interaction histories, day and night, among sampled individuals over several months to years. Social network analyses are an obvious approach to analyzing interaction data because of their flexibility for fitting many different social structures as well as the ability to assess both direct contacts and indirect associations via intermediaries. For many network properties, however, it is not clear whether estimates based upon a sample of the network are reflective of the entire network. In wildlife applications, networks may be poorly sampled and boundary effects will be common. We present an alternative approach that utilizes a hierarchical modeling framework to assess the individual, dyadic, and environmental factors contributing to variation in the interaction rates and allows us to estimate the underlying process variation in each. In a disease control context, this approach will allow managers to focus efforts on those types of individuals and environments that contribute the most toward super-spreading events. We account for the sampling distribution of proximity loggers and the non-independence of contacts among groups by only using contact data within a group during days when the group membership of proximity loggers was known. This allows us to separate the two mechanisms responsible for a pair not contacting one another: they were not in the same group or they were in the same group but did not come within the specified contact distance. We illustrate our approach with an example dataset of female elk from northwestern Wyoming and conclude with a number of important future research directions.     

Queuing for dominance: gerontocracy and queue-jumping in the hover wasp <Emphasis Type="Italic">Liostenogaster flavolineata</Emphasis>

The mechanisms through which dominance is inherited within social groups vary from direct interactions such as fighting to non-confrontational conventions. Liostenogaster flavolineata is a primitively eusocial hover wasp in which one female, the ‘dominant’, is the only reproductive upon the nest. The remaining females, although capable of reproduction, behave as helpers. In this study, we investigate the rules by which helpers inherit dominance. We removed successive dominants from 56 nests and recorded accession on un-manipulated nests. The results showed that L. flavolineata has a strict age-based inheritance queue: new dominants are the oldest female in their groups 87% of the time. Thirteen cases of queue-jumping were found in which young individuals were able to supplant older nestmates and inherit dominance precociously. Queue jumpers did not differ from other wasps in terms of relatedness to other group members or body size. Individuals that had previously worked less hard than other females of equivalent rank were significantly more likely to later jump the queue. Queue-jumping may represent a cheating strategy or could indicate that the rule for inheriting dominance is not based purely on relative age. We also discuss possible reasons why age-based queuing has evolved and its potential to promote the evolution of helping behaviour.     

Unraveling fission-fusion dynamics: how subgroup properties and dyadic interactions influence individual decisions

Many species show fission-fusion group dynamics because it has clear advantages for flexibly exploiting heterogeneous environments. However, the mechanisms by which these dynamics arise are not well known. We used a hierarchical Bayesian model to disentangle the different influences on spider monkey (Ateles geoffroyi) individual fissions and fusions, including the three dimensions of fission-fusion dynamics (subgroup size, dispersion, and composition). Furthermore, we considered the influences of other individuals also leaving or joining a subgroup at the same time. We found that the most important influence on individual fissions and fusions is whether other individuals are also doing the same. Subgroup size and dispersion did not have clear effects on the probability that an individual fissioned or fusioned, while individuals tended to leave subgroups that were biased toward the opposite sex and to join subgroups that were biased toward their own sex. The networks constructed by the inter-individual influences during fissions and fusions were cohesive and did not show assortativity by sex or by degree. Individuals had a similar degree in both networks and each was influenced by a different set of individuals, suggesting a high fluidity in the social networks. We suggest that these networks reflect the way in which information about the environment flows as individuals follow one another during fissions and fusions.     

Validation of an automated data collection method for quantifying social networks in collective behaviours

The social network of preferences among group members can affect the distribution and consequences of collective behaviours. However, the behavioural contexts and taxa in which social network structure has been described are still limited because such studies require extensive data. Here, we highlight the use of an automated passive integrated transponder (PIT)-tag monitoring system for social network analyses and do so in a novel context—nestling provisioning in an avian cooperative breeder, for which direct observation of social behaviours is difficult. First, we used observers and cameras to arrive at a suitable metric of nest visit synchrony in the PIT-tag data. Second, we validated the use of this metric for social network analyses using internal nest video cameras. Third, we used hierarchical regression models with ‘sociality’ parameter to investigate structure of networks collected from multiple groups. Use of PIT tags led to nest visitation duration and frequency being obtained with a high degree of accuracy for all group members, except for the breeding female for whom accurate estimations required the use of a video camera due to her high variability in visitation time. The PIT-tag dataset uncovered significant variability in social network structure. Our results highlight the importance of combining complementary observation methods when conducting social network analyses of wild animals. Our methods can also be generalised to multiple contexts in social systems wherever repeated encounters with other individuals in closed space have ecological implications.