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Abstract Avian mixed-species flocks are ubiquitous across habitats and a model for studying how heterospecific sociality influences the behavior and composition of animal communities. Here, we review the literature on mixed-species flocks and argue that a renewed focus on individual-level interactions among flock members can transform our understanding of this iconic, avian social system. Specifically, we suggest that an individual perspective will further our understanding of (1) how inter- and intraspecific variation in flock participation links to fitness costs and benefits, (2) the implications of familiarity between individuals in structuring mixed-species flock communities, and (3) how social roles within mixed-species flocks are related to social behavior within and across species. We summarize studies that use an individual perspective in each of these areas and discuss knowledge from conspecific social behavior to posit more broadly how individuals may shape mixed-species flocks. We encourage research approaches that incorporate individual variation in traits, relationships, and social roles in their assessment of mixed-species flocking dynamics. We propose that the analysis of individual variation in behavior will be particularly important for explicitly identifying fitness outcomes that led to the evolution of mixed-species flocks, which in turn affect community structure and resilience. 

Abstract Advances in data‐logging technologies have provided a way to monitor the movement of individual animals at unprecedented spatial and temporal scales. When used in conjunction with social network analyses, these data can provide deep insight into the structure and dynamics of animal social systems. Emergence of these new technologies demands concomitant progress in workflows to translate data streams from automated systems to social networks, based on biologically relevant metrics.Here we outline key considerations for constructing social networks from automated telemetry data. We highlight the need for paying particular attention to the spatial arrangement of receiver stations with respect to the ecology of study system and developing appropriate criteria for quantifying associations.We provide a case study for constructing social networks from automated telemetry data collected over 1 month during a study of acorn woodpeckersMelanerpes formicivorus, a cooperatively breeding bird. The data consisted of detections of known birds near receiver stations placed within core areas of group territories. We use this system to demonstrate how to build social networks to investigate biological questions about patterns of associations between group members and territory visitors across the landscape. 

Determining space use for species is fundamental to understanding their ecology, and tracking animals can reveal insights into their spatial ecology on home ranges and territories. Recent technological advances have led to GPS-tracking devices light enough for birds as small as ~30 g, creating novel opportunities to remotely monitor fine-scale movements and space use for these smaller species. We tested whether miniaturized GPS tags can allow us to understand space use of migratory birds away from their capture sites and sought to understand both pre-breeding space use as well as territory and habitat use on the breeding grounds. We used GPS tags to characterize home ranges on the breeding grounds for a migratory songbird with limited available breeding information, the Golden-crowned Sparrow (Zonotrichia atricapilla). Using GPS points from 23 individuals across 26 tags (three birds tagged twice), we found home ranges in Alaska and British Columbia were on average 44.1 ha (95% kernel density estimate). In addition, estimates of territory sizes based on field observations (mean 2.1 ha, 95% minimum convex polygon [MCP]) were three times smaller than 95% MCPs created using GPS tags (mean 6.5 ha). Home ranges included a variety of land cover classes, with shrubland particularly dominant (64–100% of home range cover for all but one bird). Three birds tracked twice returned to the same breeding area each year, supporting high breeding site fidelity for this species. We found reverse spring migration for five birds that flew up to 154 km past breeding destinations before returning. GPS-tracking technology allowed for critical ecological insights into this migratory species that breeds in very remote locations. 

Animal social interactions have an intrinsic spatial basis as many of these interactions occur in spatial proximity. This presents a dilemma when determining causality: Do individuals interact socially because they happen to share space, or do they share space because they are socially linked? We present a method that uses demographic turnover events as a natural experiment to investigate the links between social associations and space use in the context of interannual winter site fidelity in a migratory bird. We previously found that golden-crowned sparrows ( Zonotrichia atricapilla ) show consistent flocking relationships across years, and that familiarity between individuals influences the dynamics of social competition over resources. Using long-term data on winter social and spatial behavior across 10 y, we show that i) sparrows exhibit interannual fidelity to winter home ranges on the scale of tens of meters and ii) the precision of interannual site fidelity increases with the number of winters spent, but iii) this fidelity is weakened when sparrows lose close flockmates from the previous year. Furthermore, the effect of flockmate loss on site fidelity was higher for birds that had returned in more than 2 winters, suggesting that social fidelity may play an increasingly important role on spatial behavior across the lifetime of this migratory bird. Our study provides evidence that social relationships can influence site fidelity, and shows the potential of long-term studies for disentangling the relationship between social and spatial behavior. 

Cooperatively breeding species exhibit a range of social behaviours associated with different costs and benefits to group living, often in association with different environmental conditions. For example, recent phylogenetic studies have collectively shown that the evolution and distribution of cooperative breeding behaviour is related to the environment. However, little is known about how environmental variation may drive differences in social systems across populations within species, and how the relationship between environmental conditions and sociality may differ across species. Here, we examine variation in social group size along a steep environmental gradient for two congeneric cooperatively breeding species of fairywrens (Maluridae) and show that they exhibit opposing ecogeographic patterns. Purple-backed fairywrens, a species in which helpers increase group productivity, have larger groups in hot, dry environments and smaller groups in cool, wet environments. By contrast, superb fairywrens, a species with helpers that do not increase group productivity despite the presence of alloparental care, exhibit the opposite trend. We suggest differences in the costs and benefits of sociality contribute to these opposing ecogeographical patterns and demonstrate that comparisons of intraspecific patterns of social variation across species can provide insight into how ecology shapes social systems. 

Abstract Daily foraging activity of small wintering birds is classically thought to be driven by the need to gather enough energy reserves to survive each night. A separate line of research has shown that sociality is a major driver in winter foraging activities in many species. Here, we used wintering birds as a study system to move toward an integrative understanding of the influence of energy requirements and sociality on foraging ecology. We used RFID-enabled feeders in Lincoln, Nebraska, USA in January–March 2019 to measure foraging activity in two species (downy woodpeckers, Dryobates pubescens, and white-breasted nuthatches, Sitta carolinensis). We analyzed the relationship between overnight temperature and morning foraging activity and found that lowest overnight temperature was weakly correlated with morning visitation at feeders. We then used a network approach to ask if flock associations explain similarity in birds’ foraging activity. In both species, individuals with stronger associations in a social network were more likely to share similar feeder activity, and an index of social partners’ activity explained foraging activity better than overnight temperature. This brings forth new questions about the interplay between individual response to temperature and social factors in shaping how small animals cope with harsh winter conditions.