Despite growing interest in animal social networks, surprisingly little is known about whether individuals are consistent in their social network characteristics. Networks are rarely repeatedly sampled; yet an assumption of individual consistency in social behaviour is often made when drawing conclusions about the consequences of social processes and structure. A characterization of such social phenotypes is therefore vital to understanding the significance of social network structure for individual fitness outcomes, and for understanding the evolution and ecology of individual variation in social behaviour more broadly. Here, we measured foraging associations over three winters in a large PIT-tagged population of great tits, and used a range of social network metrics to quantify individual variation in social behaviour. We then examined repeatability in social behaviour over both short (week to week) and long (year to year) timescales, and investigated variation in repeatability across age and sex classes. Social behaviours were significantly repeatable across all timescales, with the highest repeatability observed in group size choice and unweighted degree, a measure of gregariousness. By conducting randomizations to control for the spatial and temporal distribution of individuals, we further show that differences in social phenotypes were not solely explained by within-population variation in local densities, but also reflected fine-scale variation in social decision making. Our results provide rare evidence of stable social phenotypes in a wild population of animals. Such stable social phenotypes can be targets of selection and may have important fitness consequences, both for individuals and for their social-foraging associates.
Both social and ecological factors influence population process and structure, with resultant consequences for phenotypic selection on individuals. Understanding the scale and relative contribution of these two factors is thus a central aim in evolutionary ecology. In this study, we develop a framework using null models to identify the social and spatial patterns that contribute to phenotypic structure in a wild population of songbirds. We used automated technologies to track 1053 individuals that formed 73 737 groups from which we inferred a social network. Our framework identified that both social and spatial drivers contributed to assortment in the network. In particular, groups had a more even sex ratio than expected and exhibited a consistent age structure that suggested local association preferences, such as preferential attachment or avoidance. By contrast, recent immigrants were spatially partitioned from locally born individuals, suggesting differential dispersal strategies by phenotype. Our results highlight how different scales of social decision-making, ranging from post-natal dispersal settlement to fission–fusion dynamics, can interact to drive phenotypic structure in animal populations.
Social network analysis has become a popular tool for characterising the social structure of populations. Animal social networks can be built either by observing individuals and defining links based on the occurrence of specific types of social interactions, or by linking individuals based on observations of physical proximity or group membership, given a certain behavioural activity. The latter approaches of discovering network structure require splitting the temporal observation stream into discrete events given an appropriate time resolution parameter. This process poses several non-trivial problems which have not received adequate attention so far. Here, using data from a study of passive integrated transponder (PIT)-tagged great tits Parus major, we discuss these problems, demonstrate how the choice of the extraction method and the temporal resolution parameter influence the appearance and properties of the retrieved network and suggest a modus operandi that minimises observer bias due to arbitrary parameter choice. Our results have important implications for all studies of social networks where associations are based on spatio-temporal proximity, and more generally for all studies where we seek to uncover the relationships amongst a population of individuals that are observed through a temporal data stream of appearance records.
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The provision of wild birds with supplementary food has increased substantially over recent decades. While it is assumed that provisioning birds is beneficial, supplementary feeding can have detrimental 'carry-over' effects on reproductive traits. Due to difficulties in monitoring individual feeding behaviour, assessing how individuals within a population vary in their exploitation of supplementary food resources has been limited. Quantifying individual consumption of supplementary food is necessary to understand the operation of carry-over effects at the individual level. We used Radio Frequency Identification (RFID) technology and automated feeders to estimate individual consumption of supplementary winter food in a large wild population of great tits Parus major and blue tits Cyanistes caeruleus. Using these data, we identified demographic factors that explained individual variation in levels of supplementary food consumption. We also tested for carry-over effects of supplementary food consumption on recruitment, reproductive success and a measure of survival. Individual variation in consumption of supplementary food was explained by differences between species, ages, sexes and years. Individuals were consistent across time in their usage of supplementary resources. We found no strong evidence that the extent of supplementary food consumption directly influenced subsequent fitness parameters. Such effects may instead result from supplementary food influencing population demographics by enhancing the survival and subsequent breeding of less competitive individuals, which reduce average breeding parameters and increase density-dependent competition. Carry-over effects of supplementary feeding are not universal and may depend upon the temporal availability of the food provided. Our study demonstrates how RFID systems can be used to examine individual-level behaviour with minimal effects on fitness.
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