Variation in local population size predicts social network structure in wild songbirds.

The structure of animal societies is a key determinant of many ecological and evolutionary processes. Yet, we know relatively little about the factors and mechanisms that underpin detailed social structure. Among other factors, social structure can be influenced by habitat configuration. By shaping animal movement decisions, heterogeneity in habitat features, such as vegetation and the availability of resources, can influence the spatiotemporal distribution of individuals and subsequently key socioecological properties such as the local population size and density. Differences in local population size and density can impact opportunities for social associations and may thus drive substantial variation in local social structure. Here, we investigated spatiotemporal variation in population size at 65 distinct locations in a small songbird, the great tit (Parus major) and its effect on social network structure. We first explored the within-location consistency of population size from weekly samples and whether the observed variation in local population size was predicted by the underlying habitat configuration. Next, we created social networks from the birds' foraging associations at each location for each week and examined if local population size affected social structure. We show that population size is highly repeatable within locations across weeks and years and that some of the observed variation in local population size was predicted by the underlying habitat, with locations closer to the forest edge having on average larger population sizes. Furthermore, we show that local population size affected social structure inferred by four global network metrics. Using simple simulations, we then reveal that much of the observed social structure is shaped by social processes. Across different population sizes, the birds' social structure was largely explained by their preference to forage in flocks. In addition, over and above effects of social foraging, social preferences between birds (i.e. social relationships) shaped certain network features such as the extent of realized social connections. Our findings thus suggest that individual social decisions substantially contribute to shaping certain social network features over and above effects of population size alone.

Social learning mechanisms shape transmission pathways through replicate local social networks of wild birds.

The emergence and spread of novel behaviours via social learning can lead to rapid population-level changes whereby the social connections between individuals shape information flow. However, behaviours can spread via different mechanisms and little is known about how information flow depends on the underlying learning rule individuals employ. Here, comparing four different learning mechanisms, we simulated behavioural spread on replicate empirical social networks of wild great tits and explored the relationship between individual sociality and the order of behavioural acquisition. Our results reveal that, for learning rules dependent on the sum and strength of social connections to informed individuals, social connectivity was related to the order of acquisition, with individuals with increased social connectivity and reduced social clustering adopting new behaviours faster. However, when behavioural adoption depends on the ratio of an individuals' social connections to informed versus uninformed individuals, social connectivity was not related to the order of acquisition. Finally, we show how specific learning mechanisms may limit behavioural spread within networks. These findings have important implications for understanding whether and how behaviours are likely to spread across social systems, the relationship between individuals' sociality and behavioural acquisition, and therefore for the costs and benefits of sociality.

Temporal avoidance as a means of reducing competition between sympatric species.

Human activity has modified the availability of natural resources and the abundance of species that rely on them, potentially changing interspecific competition dynamics. Here, we use large-scale automated data collection to quantify spatio-temporal competition among species with contrasting population trends. We focus on the spatial and temporal foraging behaviour of subordinate marsh tits Poecile palustris among groups of socially and numerically dominant blue tits Cyanistes caeruleus and great tits Parus major. The three species exploit similar food resources in mixed groups during autumn-winter. Using 421 077 winter recordings of individually marked birds at 65 automated feeding stations in Wytham Woods (Oxfordshire, UK), we found that marsh tits were less likely to join larger groups of heterospecifics, and they accessed food less frequently in larger groups than in smaller ones. Marsh tit numbers within groups declined throughout the diurnal and winter periods, while the number of blue and great tits increased. However, sites that attracted larger groups of these heterospecifics also attracted more marsh tits. The results suggest that subordinate species exhibit temporal avoidance of socially and numerically dominant heterospecifics, but have limited ability for spatial avoidance, indicating that behavioural plasticity enables only a partial reduction of interspecific competition.

Social phenotype-dependent selection of social environment in wild great and blue tits: an experimental study.

There is growing evidence that individuals actively assess the match between their phenotype and their environment when making habitat choice decisions (so-called matching habitat choice). However, to our knowledge, no studies have considered how the social environment may interact with social phenotype in determining habitat choice, despite habitat choice being an inherently social process and growing evidence for individual variation in sociability. We conducted an experiment using wild great and blue tits to understand how birds integrate their social phenotype and social environment when choosing where and how to feed. We used programmable feeders to (i) record social interactions and estimate social phenotype, and (ii) experimentally manipulate the local density experienced by birds of differing social phenotype. By tracking feeder usage, we estimated how social environment and social phenotype predicted feeder choice and feeding behaviour. Both social environment and social phenotype predicted feeder usage, but a bird's decision to remain in a particular social environment did not depend on their social phenotype. By contrast, for feeding behaviour, responses to the social environment depended on social phenotype. Our results provide rare evidence of matching habitat choice and shed light on the dependence of habitat choice on between-individual differences in social phenotype.

Social network position predicts male mating success in a small passerine.

Individuals differ in the quantity and quality of their associations with conspecifics. The resulting variation in the positions that individuals occupy within their social environment can affect several aspects of life history, including reproduction. While research increasingly shows how social factors can predict dyadic mating patterns (who will breed with whom), much less is known about how an individual's social position affects its overall likelihood to acquire mating partner(s). We studied social networks of socially monogamous blue tits (Cyanistes caeruleus) to investigate whether the number and strength of connections to opposite-sex conspecifics, the ratio between same- and opposite-sex connections, and the tendency to move between social groups in the months prior to breeding affect individuals' success in acquiring 1) a breeding partner and 2) an extrapair partner. After controlling for differences in spatial location, we show that males that moved more often between social groups were more likely to acquire a breeding partner. Moreover, adult males that associated with more females were more likely to sire extrapair young. The number of female associates also predicted the proportion of familiar female breeding neighbors, suggesting that familiarity among neighbors may facilitate opportunities for extrapair matings. In females, none of the network metrics significantly predicted the likelihood of acquiring a breeding or extrapair partner. Our study suggests that the positioning of males within their social environment prior to breeding can translate into future mating success, adding an important new dimension to studies of (extrapair) mating behavior.

Animal behavior: Innovation in the city.

Behavioral innovations may help animals cope with new environments, but how such behaviors start is hard to capture. A new study reports the innovation and transmission of a new foraging culture in an urban parrot.

The importance of individual-to-society feedbacks in animal ecology and evolution.

The social decisions that individuals make-who to interact with and how frequently-give rise to social structure. The resulting social structure then determines how individuals interact with their surroundings-resources and risks, pathogens and predators, competitors and cooperators. However, despite intensive research on (a) how individuals make social decisions and (b) how social structure shapes social processes (e.g. cooperation, competition and conflict), there are still few studies linking these two perspectives. These perspectives represent two halves of a feedback loop: individual behaviour scales up to define the social environment, and this environment, in turn, feeds back by shaping the selective agents that drive individual behaviour. We first review well-established research areas that have captured both elements of this feedback loop-host-pathogen dynamics and cultural transmission. We then highlight areas where social structure is well studied but the two perspectives remain largely disconnected. Finally, we synthesise existing research on 14 distinct research topics to identify new prospects where the interplay between social structure and social processes are likely to be important but remain largely unexplored. Our review shows that the inherent links between individuals' traits, their social decisions, social structure and social evolution, warrant more consideration. By mapping the existing and missing connections among many research areas, our review highlights where explicitly considering social structure and the individual-to-society feedbacks can reveal new dimensions to old questions in ecology and evolution.

Analysis of within-individual variation in extrapair paternity in blue tits (Cyanistes caeruleus) shows low repeatability and little effect of changes in neighborhood.

Many studies investigated variation in the frequency of extrapair paternity (EPP) among individuals. However, our understanding of within-individual variation in EPP remains limited. Here, we comprehensively investigate variation in EPP at the within-individual level in a population of blue tits (Cyanistes caeruleus). Our study is based on parentage data comprising >10 000 genotyped offspring across 11 breeding seasons. First, we examined the repeatability of the occurrence of EPP, the number of extrapair offspring, the number of extrapair partners, and the occurrence of paternity loss using data from males and females that bred in multiple years. Second, we tested whether within-individual changes in EPP between breeding seasons relate to between-year changes in the local social environment. Repeatabilities were generally low but significant for the occurrence and number of extrapair young in females and for whether a male sired extrapair young or not. We found no evidence that the presence of the former social partner or changes in the proportion of familiar individuals or in phenotypic traits of the neighbors influenced changes in levels of EPP in females. However, in adult males, a decrease in the average body size of male neighbors was associated with higher extrapair siring success. If confirmed, this result suggests that the competitive ability of a male relative to its neighbors influences his extrapair mating success. We suggest that alternative hypotheses, including the idea that within-individual changes in EPP are due to "chance events" rather than changes in an individual's social breeding environment, deserve more consideration.

Winter associations predict social and extra-pair mating patterns in a wild songbird.

Despite decades of research, our understanding of the underlying causes of within-population variation in patterns of extra-pair paternity (EPP) remains limited. Previous studies have shown that extra-pair mating decisions are linked to both individual traits and ecological factors. Here, we examine whether social associations among individuals prior to breeding also shape mating patterns, specifically the occurrence of EPP, in a small songbird, the blue tit. We test whether associations during the non-breeding period predict (1) future social pairs, (2) breeding proximity (i.e. the distance between breeding individuals) and (3) the likelihood that individuals have extra-pair young together. Individuals that were more strongly associated (those that foraged more often together) during winter tended to nest closer together. This, by itself, predicts EPP patterns, because most extra-pair sires are close neighbours. However, even after controlling for spatial effects, female-male dyads with stronger social associations prior to breeding were more likely to have extra-pair young. Our findings reveal a carry-over from social associations into future mating decisions. Quantifying the long-term social environment of individuals and studying its dynamics is a promising approach to enhance our understanding of the process of (extra-)pair formation.

Relying on known or exploring for new? Movement patterns and reproductive resource use in a tadpole-transporting frog.

Animals relying on uncertain, ephemeral and patchy resources have to regularly update their information about profitable sites. For many tropical amphibians, widespread, scattered breeding pools constitute such fluctuating resources. Among tropical amphibians, poison frogs (Dendrobatidae) exhibit some of the most complex spatial and parental behaviors-including territoriality and tadpole transport from terrestrial clutches to ephemeral aquatic deposition sites. Recent studies have revealed that poison frogs rely on spatial memory to successfully navigate through their environment. This raises the question of when and how these frogs gain information about the area and suitable reproductive resources. To investigate the spatial patterns of pool use and to reveal potential explorative behavior, we used telemetry to follow males of the territorial dendrobatid frog Allobates femoralis during tadpole transport and subsequent homing. To elicit exploration, we reduced resource availability experimentally by simulating desiccated deposition sites. We found that tadpole transport is strongly directed towards known deposition sites and that frogs take similar direct paths when returning to their home territory. Frogs move faster during tadpole transport than when homing after the deposition, which probably reflects different risks and costs during these two movement phases. We found no evidence for exploration, neither during transport nor homing, and independent of the availability of deposition sites. We suggest that prospecting during tadpole transport is too risky for the transported offspring as well as for the transporting male. Relying on spatial memory of multiple previously discovered pools appears to be the predominant and successful strategy for the exploitation of reproductive resources in A. femoralis. Our study provides for the first time a detailed description of poison frog movement patterns during tadpole transport and corroborates recent findings on the significance of spatial memory in poison frogs. When these frogs explore and discover new reproductive resources remains unknown.