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.
Social and genetic mating systems play an important role in natural and sexual selection, as well as in the dynamics of populations. In socially monogamous species different genetic mating patterns appear when individuals mate outside the breeding pair within a breeding season (extra-pair mating) or when they change partners between two breeding seasons (widowing or divorce). Divorce can be defined as having occurred when two previously paired individuals are alive during the next breeding season and at least one of them has re-mated with a new partner. In socially monogamous birds divorce is widespread, but it is not clear whether it is a behavioural adaptation to improve the quality of a mating decision or whether, alternatively, it results as a non-selected consequence of other processes: existing studies suggest a heterogeneous set of results with respect to this central question. This heterogeneity could result from a number of factors, ranging from the methodological approaches used, to population- or species-specific characters. In this review we use phylogenetic meta-analyses to assess the evidence that divorce is adaptive (in terms of breeding success) across 64 species of socially monogamous birds. Second, we explore biological and methodological reasons for the heterogeneity in the results of previous studies. Results of our analyses supported the hypothesis that divorce is, in general, an adaptive behavioural strategy as: (1) divorce is triggered by relatively low breeding success; (2) there is a positive change in breeding success as a result of divorce. More specifically, while controlling for methodological moderators, we show that: (i) earlier stages of breeding are better predictors of divorce than later stages (r = 0.231; 95% CI: 0.061-0.391 for clutch size; similar for laying date); (ii) females benefited from divorce more than males in terms of increasing breeding success between successive breeding attempts, with different stages of the breeding cycle improving at different rates (e.g. r = 0.637; 95% CI: 0.328-0.817 for brood-level measures). We show that the effect size was dependent on the methodological approach used across studies and argue that research on the adaptive nature of divorce should be cautious when designing the study and interpreting the results. Altogether, by providing strong evidence that divorce is an adaptive strategy across monogamous birds, the results of our analysis provide a firm ground for further exploration of external covariates of divorce (e.g. demographic factors) and the mechanisms underlying the differences in the effect sizes of the proximal fitness causes and consequences of divorce.
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.
Access to analytical code is essential for transparent and reproducible research. We review the state of code availability in ecology using a random sample of 346 nonmolecular articles published between 2015 and 2019 under mandatory or encouraged code-sharing policies. Our results call for urgent action to increase code availability: only 27% of eligible articles were accompanied by code. In contrast, data were available for 79% of eligible articles, highlighting that code availability is an important limiting factor for computational reproducibility in ecology. Although the percentage of ecological journals with mandatory or encouraged code-sharing policies has increased considerably, from 15% in 2015 to 75% in 2020, our results show that code-sharing policies are not adhered to by most authors. We hope these results will encourage journals, institutions, funding agencies, and researchers to address this alarming situation.
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