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.     

Spatial and temporal heterogeneities in the contact behaviour of rabbits

It is becoming increasingly evident that the social behaviour of many group-living species is more complex than previously assumed and that free mixing of individuals, even within social groups, is rare. This has important implications for ecological processes, such as disease transmission, which are dependent on interactions between individuals. European wild rabbits (Oryctolagus cuniculus) have been considered traditionally as highly sociable animals that mix freely within groups but interact less frequently between groups. We deployed proximity logging devices to quantify the intra- and inter-group contact behaviour of free-living wild rabbits in two populations in a temperate region of Australia. Altogether, 126 rabbits were fitted with proximity loggers at least once during the study. Radio-tracking was carried out alongside proximity data collection to determine the space use of rabbit social groups within the study sites. On average, a rabbit made only 1.54 ± 0.23 (SE) (median = 0.54) contacts per day with each other rabbit carrying a proximity logger in its social group, and the mean daily contact duration was 202 ± 38 s/day (SE) (median = 29 s). Despite the high degree of home range overlap between the neighbouring social groups, inter-group contacts were highly infrequent and brief. Our results demonstrated considerable spatial and temporal heterogeneities in the contact behaviour between individual rabbits, both between populations and between and within social groups in the same population. Such variations in the social organisation of rabbits are likely to create complex patterns of disease transmission through direct contact and may contribute towards observed heterogeneities in the effects of disease on wild rabbit populations.     

Spatial,socio‐economic,and ecological implications of incorporating minimum size constraints in marine protected area network design

Marine protected areas (MPAs) are the cornerstone of most marine conservation strategies, but the effectiveness of each one partly depends on its size and distance to other MPAs in a network. Despite this, current recommendations on ideal MPA size and spacing vary widely, and data are lacking on how these constraints might influence the overall spatial characteristics, socio‐economic impacts, and connectivity of the resultant MPA networks. To address this problem, we tested the impact of applying different MPA size constraints in English waters. We used the Marxan spatial prioritization software to identify a network of MPAs that met conservation feature targets, whilst minimizing impacts on fisheries; modified the Marxan outputs with the MinPatch software to ensure each MPA met a minimum size; and used existing data on the dispersal distances of a range of species found in English waters to investigate the likely impacts of such spatial constraints on the region's biodiversity. Increasing MPA size had little effect on total network area or the location of priority areas, but as MPA size increased, fishing opportunity cost to stakeholders increased. In addition, as MPA size increased, the number of closely connected sets of MPAs in networks and the average distance between neighboring MPAs decreased, which consequently increased the proportion of the planning region that was isolated from all MPAs. These results suggest networks containing large MPAs would be more viable for the majority of the region's species that have small dispersal distances, but dispersal between MPA sets and spill‐over of individuals into unprotected areas would be reduced. These findings highlight the importance of testing the impact of applying different MPA size constraints because there are clear trade‐offs that result from the interaction of size, number, and distribution of MPAs in a network.     

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.     

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.     

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.     

Seasonal changes in the structure of rhesus macaque social networks

Social structure emerges from the patterning of interactions between individuals and plays a critical role in shaping some of the main characteristics of animal populations. The topological features of social structure, such as the extent to which individuals interact in clusters, can influence many biologically important factors, including the persistence of cooperation, and the rate of spread of disease. Yet, the extent to which social structure topology fluctuates over relatively short periods of time in relation to social, demographic, or environmental events remains unclear. Here, we use social network analysis to examine seasonal changes in the topology of social structures that emerge from socio-positive associations in adult female rhesus macaques (Macaca mulatta). Behavioral data for two different association types (grooming and spatial proximity) were collected for females in two free-ranging groups during two seasons: the mating and birth seasons. Stronger dyadic bonds resulted in social structures that were more tightly connected (i.e., of greater density) in the mating season compared to the birth season. Social structures were also more centralized around a subset of individuals and more clustered in the mating season than those in the birth season, although the latter differences were mostly driven by differences in density alone. Our results suggest a degree of temporal variation in the topological features of social structure in this population. Such variation may feed back on interactions, hence affecting the behaviors of individuals, and may therefore be important to take into account in studies of animal behavior.     

I know you: familiarity with an audience influences male-male interactions in Siamese fighting fish, Betta splendens

The presence of an audience alters male–male interactions in a number of social species. Additionally, whether individuals have interacted previously may influence the current encounter. Although it is clear that previous exposure influences communication and that the presence of an audience affects behavior in a number of contexts, how these phenomena might interact to impact interactions is underexplored. This study examined the effects of audience familiarity on male–male interactions in Siamese fighting fish (Betta splendens). Treatments that varied based on familiarity with and sex of the audience as well as length of time after initial audience exposure were used to investigate whether male aggressive interactions were affected by these factors. Males first encountered an individual that would later become an audience. Pairs of males then interacted with each other in either the presence of this familiar individual or a new, unfamiliar audience conspecific or with no audience at all. Opponent-directed gill flaring and tail beats were influenced by familiarity with the audience individual, although they were affected differently by the sex of the audience present. Gill flaring was affected predominantly by the presence of a male audience, whereas tail beat rate was influenced by the presence of a female audience. Gill flaring rate did not differ regardless of whether 4 or 24?h had passed between the initial exposure to the audience and the testing phase. In contrast, the familiarity effect disappeared after 4?h for tail beats. These results stress the importance of considering the familiarity of individuals within a network when examining communication networks.     

The influence of refuge sharing on social behaviour in the lizard Tiliqua rugosa

Refuge sharing by otherwise solitary individuals during periods of inactivity is an integral part of social behaviour and has been suggested to be the precursor to more complex social behaviour. We compared social association patterns of active versus inactive sheltering individuals in the social Australian sleepy lizard, Tiliqua rugosa, to empirically test the hypothesis that refuge sharing facilitates social associations while individuals are active. We fitted 18 neighbouring lizards with Global Positioning System (GPS) recorders to continuously monitor social associations among all individuals, based on location records taken every 10 min for 3 months. Based on these spatial data, we constructed three weighted, undirected social networks. Two networks were based on empirical association data (one for active and one for inactive lizards in their refuges), and a third null model network was based on hypothetical random refuge sharing. We found patterns opposite to the predictions of our hypothesis. Most importantly, association strength was higher in active than in inactive sheltering lizards. That is, individual lizards were more likely to associate with other lizards while active than while inactive and in shelters. Thus, refuge sharing did not lead to increased frequencies of social associations while lizards were active, and we did not find any evidence that refuge sharing was a precursor to sleepy lizard social behaviour. Our study of an unusually social reptile provides both quantitative data on the relationship between refuge sharing and social associations during periods of activity and further insights into the evolution of social behaviour in vertebrates.     

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.     

Determining association networks in social animals: choosing spatial–temporal criteria and sampling rates

Social Network Analysis has become an important methodological tool for advancing our understanding of human and animal group behaviour. However, researchers tend to rely on arbitrary distance and time measures when defining ‘contacts’ or ‘associations’ between individuals based on preliminary observation. Otherwise, criteria are chosen on the basis of the communication range of sensor devices (e.g. bluetooth communication ranges) or the sampling frequencies of collection devices (e.g. Global Positioning System devices). Thus, researchers lack an established protocol for determining both relevant association distances and minimum sampling rates required to accurately represent the network structure under investigation. In this paper, we demonstrate how researchers can use experimental and statistical methods to establish spatial and temporal association patterns and thus correctly characterise social networks in both time and space. To do this, we first perform a mixing experiment with Merino sheep (Ovis aries) and use a community detection algorithm that allows us to identify the spatial and temporal distance at which we can best identify clusters of previously familiar sheep. This turns out to be within 2–3 m of each other for at least 3 min. We then calculate the network graph entropy rate—a measure of ease of spreading of information (e.g. a disease) in a network—to determine the minimum sampling rate required to capture the variability observed in our sheep networks during distinct activity phases. Our results indicate the need for sampling intervals of less than a minute apart. The tools that we employ are versatile and could be applied to a wide range of species and social network datasets, thus allowing an increase in both the accuracy and efficiency of data collection when exploring spatial association patterns in gregarious species.     

Ontogenetic changes in alarm-call production and usage in meerkats (<Emphasis Type="Italic">Suricata suricatta</Emphasis>): adaptations or constraints?

In many species, individuals suffer major mortality in their first year because of predation. Behaviours that facilitate successful escape are therefore under strong selection, but anti-predator skills often emerge gradually during an individual’s early development. Using long-term data and acoustic recordings of alarm calls collected during natural predator encounters, we aimed to elucidate two largely unsolved issues in anti-predator ontogeny: (1) whether incorrect predator assignment is adaptively age-appropriate, given that vulnerability often changes during development, or whether age-related differences reflect true mistakes made by immature individuals; and (2) the extent to which the development of adult-like competence in alarm-call production and usage is simply a function of maturational processes or dependent upon experience. We found that young meerkats (Suricata suricatta) were less likely to give alarm calls than adults, but alarmed more in response to non-threatening species compared to adults. However, stimuli that pose a greater threat to young than adults did not elicit more calling from young; this argues against age-related changes in vulnerability as the sole explanation for developmental changes in calling. Young in small groups, who were more likely to watch out for predators, alarmed more than less vigilant young in larger groups. Moreover, despite similarities in acoustic structure between alarm call types, calls appeared in the repertoire at different rates, and those that were associated with frequently encountered predators were produced relatively early on. These results indicate that experience is a more plausible explanation for such developmental trajectories than maturation.     

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.     

Olfactory discrimination of age-specific hydrocarbons generates behavioral segregation in a honeybee colony

The functioning of a honeybee colony relies on the coordination of colony activities via inter-individual interactions. While the structure of this interaction network keeps the young individuals relatively isolated from the rest of the colony, there are two possible mechanisms that can generate this organizational immunity. A spatial segregation that restricts the young bees to the center of the colony can shield them with equal effectiveness as a behavioral segregation in which old bees choose to interact with young bees less frequently. We test the role of these two mechanisms by determining the interaction frequency between different age groups and testing their correlation with the olfactory sensitivity of different age groups to the cuticular odor of each other. Young bees were found to interact with bees of all age groups with equal frequency, which correlates with their lack of olfactory bias for any specific age, while old bees interacted more with other old bees, which correlates with their higher olfactory sensitivity toward the cuticular odor of old bees. The distribution of olfactory responsiveness was found to be positively skewed for old bees, which provides a mechanistic basis for the heterogeneous connectivity of the interaction network observed in an earlier study. As old bees are more likely to be responsible for introducing a potential disease into the colony from the outside and spreading it via the interaction network, these results suggest that behavioral segregation, mediated by olfactory discrimination, plays an important role in generating the organizational immunity within the honeybee colony.     

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.     

Replacement of breeders and within-group conflict in the cooperatively breeding acorn woodpecker

Summary Breeding males or females were removed from groups of the cooperatively breeding acorn woodpecker to study (1) competition among individuals from different groups to fill the breeding vancancies, (2) conflict between parents and offspring within groups over replacement of the removed bird, and (3) conflict between parents and offspring for breeding status. Intense fights (power struggles) among nonbreeders from other groups to fill the breeding vacancies developed after the removals. In groups containing only helpers of the opposite sex as the removed bird, replacement was rapid, but in groups containing helpers of the same sex as the removed bird replacement was delayed substantially. In these latter groups parent-offspring conflict developed: helpers could not breed because their parent of the opposite sex was present, and they attacked potential replacements, while their remaining parent advertised for a new mate by calling. Once replacement had occurred, dominance interactions between parents and offspring and among siblings appeared to prevent some individuals from breeding. We conclude that the potential for competition among close kin is high in cooperative species because breeding opportunities are limited.     

Determining the community structure of the coral Seriatopora hystrix from hydrodynamic and genetic networks

The exchange of genetic information between coral reefs through the transport of larvae can be described in terms of networks that capture the linkages between distant populations. A key question arising from these networks is the determination of the highly connected modules (communities). Communities can be defined using genetic similarity or distance statistics between multiple samples but due to limited specimen sampling capacity the boundaries of the communities for the known coral reefs in the seascape remain unresolved. In this study we use the microsatellite composition of individual corals to compare sample populations using a genetic dissimilarity measure (F_ST) which is then used to create a complex network. This network involved sampling 1025 colonies from 22 collection sites and examining 10 microsatellites loci. The links between each sampling site were given a strength that was created from the pair wise F_ST values. The result is an undirected weighted network describing the genetic dissimilarity between each sampled population. From this network we then determined the community structure using a leading eigenvector algorithm within graph theory. However, given the relatively limited sampling conducted, the representation of the regional genetic structure was incomplete. To assist with defining the boundaries of the genetically based communities we also integrated the communities derived from a hydrodynamic and distance based networks. The hydrodynamic network, though more comprehensive, was of smaller spatial extent than our genetic sampling. A Bayesian Belief network was developed to integrate the overlapping communities. The results indicate the genetic population structure of the Great Barrier Reef and provide guidance on where future genetic sampling should take place to complete the genetic diversity mapping.     

Identifying important species that amplify or mitigate the interactive effects of human impacts on marine food webs

Some species may have a larger role than others in the transfer of complex effects of multiple human stressors, such as changes in biomass, through marine food webs. We devised a novel approach to identify such species. We constructed annual interaction-effect networks (IENs) of the simulated changes in biomass between species of the southeastern Australian marine system. Each annual IEN was composed of the species linked by either an additive (sum of the individual stressor response), synergistic (lower biomass compared with additive effects), or antagonistic (greater biomass compared with additive effects) response to the interaction effect of ocean warming, ocean acidification, and fisheries. Structurally, over the simulation period, the number of species and links in the synergistic IENs increased and the network structure became more stable. The stability of the antagonistic IENs decreased and became more vulnerable to the loss of species. In contrast, there was no change in the structural attributes of species linked by an additive response. Using indices common in food-web and network theory, we identified the species in each IEN for which a change in biomass from stressor effects would disproportionately affect the biomass of other species via direct and indirect local, intermediate, and global predator–prey feeding interactions. Knowing the species that transfer the most synergistic or antagonistic responses in a food-web may inform conservation under increasing multiple-stressor impacts.