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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 preferen

Major climatic changes in conjunction with animal movement may be associated with the spread of parasites and their vectors into new populations, with potentially important consequences for population persistence. Parasites can evolve to adapt to unsuitable ecological conditions and take up refuge within new host species, with consequences for the population growth of the new host species. One parasite species that has likely been increasing its geographic range, and potentially infecting new hosts, is the recently described air sac nematode Serratospiculoides amaculata, in great tits (Parus major) in Slovakia. In this study, we screened wild birds for potential air sac nematode infection in a woodland area of southern Germany. We identified four additional host species: Eurasian nuthatch, great spotted woodpecker, greenfinch and robin. As infection by this group of nematodes can be highly pathogenic, we recommend further investigation into its potential risk to these populations.

Individuals show consistent between-individual behavioural variation when they interact with conspecifics or heterospecifics. Such patterns might underlie emergent group-specific behavioural patterns and between-group behavioural differences. However, little is known about (i) how social and non-social drivers (external drivers) shape group-level social structures and (ii) whether animal groups show consistent between-group differences in social structure after accounting for external drivers. We used automated tracking to quantify daily social interactions and association networks in 12 colonies of zebra finches (Taeniopygia guttata). We quantified the effects of five external drivers (group size, group composition, ecological factors, physical environments and methodological differences) on daily interaction and association networks and tested whether colonies expressed consistent differences in day-to-day network structure after controlling for these drivers. Overall, we found that external drivers contribute significantly to network structure. However, even after accounting for the contribution of external drivers, there remained significant support for consistent between-group differences in both interaction (repeatability R: up to 0.493) and association (repeatability R : up to 0.736) network structures. Our study demonstrates how group-level differences in social behaviour can be partitioned into different drivers of variation, with consistent contributions from both social and non-social factors.

Multilevel societies are formed when stable groups of individuals spatially overlap and associate preferentially with other groups, producing a hierarchical social structure1. Once thought to be exclusive to humans and large mammals, these complex societies have recently been described in birds2,3. However, it remains largely unclear what benefits individuals gain by forming multilevel societies1. One hypothesis—based on food sharing in hunter-gatherers4—is that multilevel societies facilitate access to a range of cooperative relationships, with individual investment varying across the hierarchical levels of the society. We tested experimentally if such graded cooperation occurs in the multilevel society of a songbird, the superb fairy-wren (Malurus cyaneus). Specifically, we measured whether responses to playbacks of distress calls—used to recruit help when in extreme danger—varied according to the social level at which the focal individual is connected with the caller. We predicted that anti-predator responses should be highest within breeding groups (the core social unit), intermediate between groups from the same community, and lowest across groups from different communities. Our results confirm that birds exhibit the predicted hierarchical pattern of helping, and that, within breeding groups, this pattern was independent of kinship. This pattern of graded helping responses supports the hypothesis that multilevel social structures can sustain stratified cooperative relationships, and reveals similarity in cooperation in qualitatively different behaviours–anti-predator behaviour and food sharing–in the multilevel societies of songbirds and humans.

How individuals’ prior experience and population evolutionary history shape emergent patterns in animal collectives remains a major gap in the study of collective behaviour. One reason for this is that the processes that can shape individual contributions to collective actions can happen over very different timescales from each other and from the collective actions themselves, resulting in mismatched timescales. For example, a preference to move towards a specific patch might arise from phenotype, memory or physiological state. Although providing critical context to collective actions, bridging different timescales remains conceptually and methodologically challenging. Here, we briefly outline some of these challenges, and discuss existing approaches that have already generated insights into the factors shaping individual contributions in animal collectives. We then explore a case study of mismatching timescales—defining relevant group membership—by combining fine-scaled GPS tracking data and daily field census data from a wild population of vulturine guineafowl ( Acryllium vulturinum ). We show that applying different temporal definitions can produce different assignments of individuals into groups. These assignments can then have consequences when determining individuals' social history, and thus the conclusions we might draw on the impacts of the social environment on collective actions. This article is part of a discussion meeting issue ‘Collective behaviour through time’.

Spatial and social behaviour are fundamental aspects of an animal's biology, and their social and spatial environments are indelibly linked through mutual causes and shared consequences. We define the 'spatial-social interface' as intersection of social and spatial aspects of individuals' phenotypes and environments. Behavioural variation at the spatial-social interface has implications for ecological and evolutionary processes including pathogen transmission, population dynamics, and the evolution of social systems. We link spatial and social processes through a foundation of shared theory, vocabulary, and methods. We provide examples and future directions for the integration of spatial and social behaviour and environments. We introduce key concepts and approaches that either implicitly or explicitly integrate social and spatial processes, for example, graph theory, density-dependent habitat selection, and niche specialization. Finally, we discuss how movement ecology helps link the spatial-social interface. Our review integrates social and spatial behavioural ecology and identifies testable hypotheses at the spatial-social interface.

GPS-based tracking is widely used for studying wild social animals. Much like traditional observational methods, using GPS devices requires making a number of decisions about sampling that can affect the robustness of a study's conclusions. For example, sampling fewer individuals per group across more distinct social groups may not be sufficient to infer group- or subgroup-level behaviours, while sampling more individuals per group across fewer groups limits the ability to draw conclusions about populations. Here, we provide quantitative recommendations when designing GPS-based tracking studies of animal societies. We focus on the trade-offs between three fundamental axes of sampling effort: (1) sampling coverage—the number and allocation of GPS devices among individuals in one or more social groups; (2) sampling duration—the total amount of time over which devices collect data and (3) sampling frequency—the temporal resolution at which GPS devices record data. We first test GPS tags under field conditions to quantify how these aspects of sampling design can affect both GPS accuracy (error in absolute positional estimates) and GPS precision (error in the estimate relative position of two individuals), demonstrating that GPS error can have profound effects when inferring distances between individuals. We then use data from whole-group tracked vulturine guineafowl Acryllium vulturinum to demonstrate how the trade-off between sampling frequency and sampling duration can impact inferences of social interactions and to quantify how sampling coverage can affect common measures of social behaviour in animal groups, identifying which types of measures are more or less robust to lower coverage of individuals. Finally, we use data-informed simulations to extend insights across groups of different sizes and cohesiveness. Based on our results, we are able to offer a range of recommendations on GPS sampling strategies to address research questions across social organizational sc