Drivers of passive leadership in wild songbirds: species-level differences and spatio-temporally dependent intraspecific effects

Collective behaviors are typical for many social species and can have fitness benefits for participating individuals. To maximize the benefits obtained from group living, individuals must coordinate their behaviors to some extent. What are the mechanisms that make certain individuals more likely to initiate collective behaviors, for example, by taking a risk to initially access a resource (i.e., to act as “leaders”)? Here, we examine leading behavior in a natural population of great tits and blue tits. We use automated feeding stations to monitor the feeder visits of tagged individuals within mixed-species flocks, with a small cost (waiting < 2 s) associated with the initial unlocking of the feeder. We find that great tits, males, and individuals with high activity levels were more likely to be leading in each of their feeder visits. Using a null model approach, we demonstrate that the effects of sex and activity on passive leading behavior can be explained by patterns of spatial and temporal occurrence. In other words, these effects can be explained by the times and locations of when individuals visit rather than the actual order of arrival. Hence, an analysis of the causes of leading behavior is needed to separate the effects of different processes. We highlight the importance of understanding the mechanisms behind leading behavior and discuss directions for future experimental work to gain a better understanding of the causes of leadership in natural populations. Significance statement Many species are social and engage in collective behaviors. To benefit from group actions, individuals need to fulfill different roles. Here, we examine leading behavior during feeding events; who feeds first when birds arrive at a resource? In mixed-species flocks of passerines, great tits (the larger and more dominant species), males, and individuals with higher levels of activity lead more often than blue tits, females, and individuals with lower levels of activity. While the species effect remains even when we control for the locations and dates of individual feeder visits, the effects of sex and activity are dependent on when and where birds choose to feed.

The composition of mixed-species flocks of birds in and around Chitwan National Park, Nepal

Background Mixed-species flocks (MSFs) have been well sampled in the South Asia, but there has been as yet surprisingly little work on MSFs of Nepal, despite a diverse and well-studied avifauna. We surveyed MSFs in two forest types in and around the Important Bird Area of Chitwan National Park in Nepal, between 150 and 800 m a.s.l., to provide a first description of the composition of MSFs in this area. We also aimed to understand which species should be considered ‘nuclear species’, important to forming MSFs or leading them forward. Results In total, we collected records on 222 MSFs that included 100 species, and 6097 individuals. The MSFs were similar to worldwide patterns in being dominated by leaf-gleaning, non-terrestrial insectivores. However, the MSFs were more dominated by canopy species than usual, and did not have a clear gregarious, understory leading species. Rather drongos (Family Dicruridae) and minivets (Family Campephagidae, Genus Pericrocotus) acted as leaders, and a cluster analysis of composition showed one group of large body size MSFs particularly characterized by the presence of the Greater Racket-tailed Drongo (Dicrurus paradiseus). Conclusions Drongos are known to provide both costs and benefits to other flock participants: they are aggressive birds that can steal food, and manipulate other species with their vocalizations, but at the same time they are ‘sentinel species’ that produce information about predation risk other species can use. This study demonstrates that drongos can be considered nuclear species for some types of MSFs, despite the potential costs of their presence. MSFs led by sentinel species thus may form in Asia, as well as in the Neotropics.

Context-specific behavior serves as a mechanism of interspecific cohesion in mixed-species flocks

We investigated whether context-specific behavior is responsible for the cohesion of mixed-species flocks of antshrikes and antwrens in Amazonian Peru. Antshrikes perform a behavior while spatially repositioning, to which antwrens respond by approaching. The cohesion of interspecific associations requires communication, although the mechanisms often are unexplored. In monospecific groups, cohesion among individuals is maintained with actions or vocalizations given in a certain context. Dusky-throated Antshrikes (Thamnomanes ardesiacus) vocalize while in flight and the number of times they vocalize covaries with the flight distance. We refer to this pairing of flight and vocalization as repositioning behavior. Furthermore, antshrikes pair a different call type with perching, which we refer to as perching behavior. We followed Long-winged (Myrmotherula longipennis) and White-flanked Antwrens (M. axillaris) and recorded their response following natural vocalizations (no playback used) given by the antshrikes. Long-winged Antwrens, but not White-flanked, flew toward an antshrike significantly sooner and were more likely to approach the antshrikes after the repositioning behavior than after perching behavior. In addition, Long-winged Antwrens, but not White-flanked, flew toward an antshrike sooner after a longer series of repositioning calls than after a shorter series. We did not distinguish between the Long-winged Antwrens’ response as a function of movement vs. vocalizations of the antshrikes, although we argue that vocalizations are likely a more important communication component of repositioning behavior than movement. It remains unclear whether the antshrikes are deliberately signaling the Long-winged Antwrens or the antwrens are taking advantage of the repositioning behavior; active signaling is possible since antshrikes benefit from antwrens. White-flanked Antwrens may be less responsive to the antshrikes since they have a lower propensity to occur in a mixed-species flock. The results indicate that the repositioning behavior of Dusky-throated Antshrikes is a key mechanism of interspecific cohesion of Amazonian mixed-species flocks of the understory.

Temperature and vegetation complexity structure mixed-species flocks along a gradient of elevation in the tropical Andes

Mixed-species flocks constitute community modules that can help test mechanisms driving changes to community composition across environmental gradients. Here, we examined elevational patterns of flock diversity (species richness, taxonomic diversity, species, and guild composition) and asked if these patterns were reflections of the full bird community at a given elevation (open-membership hypothesis), or if they were instead structured by environmental variables. We surveyed both the overall avian community and mixed-species flocks across an undisturbed elevational gradient (~1,350–3,550 m) in the Bolivian Andes. We then tested for the role of temperature (a surrogate for abiotic stress), resource diversity (arthropods, fruits), and foraging niche diversity (vegetation vertical complexity) in structuring these patterns. Patterns for the overall and flocking communities were similar, supporting our open-membership hypothesis that Andean flocks represent dynamic, unstructured aggregations. Membership openness and the resulting flock composition, however, also varied with elevation in response to temperature and vegetation complexity. We found a mid-elevation peak in flock species richness, size, and Shannon’s diversity at ~2,300 m. The transition of flocking behavior toward a more open-membership system at this elevation may explain a similar peak in the proportion of insectivores joining flocks. At high elevations, increasing abiotic stress and decreasing fruit diversity led more generalist, gregarious tanagers (Thraupidae) to join flocks, resulting in larger yet more even flocks alongside a loss of vegetation structure. At lower elevations, flock species richness increased with greater vegetation complexity, but a greater diversity of foraging niches resulted in flocks that were more segregated into separate canopy and understory sub-types. This segregation likely results from increased costs of interspecific competition and activity matching (i.e., constraints on movement and foraging rate) for insectivores. Mid-elevation flocks (~2,300 m) seemed, therefore, to benefit from both the open-membership composition of high-elevation flocks and the high vegetation complexity of mid- and low-elevation forests.

Phenotypic Clumping Decreases With Flock Richness in Mixed-Species Bird Flocks

Animals that live in groups may experience positive interactions such as cooperative behavior or negative interactions such as competition from group members depending on group size and similarity between individuals. The effect of group size and phenotypic and ecological similarity on group assembly has not been well-studied. Mixed-species flocks are important subsets of bird communities worldwide. We examined associations within these in relation to flock size, to understand rules of flock assembly, in the Western Ghats of India. We examined the relationship between phenotypic clumping and flock richness using four variables—body size, foraging behavior, foraging height and taxonomic relatedness. Using a null model approach, we found that small flocks were more phenotypically clumped for body size than expected by chance; however, phenotypic clumping decreased as flocks increased in size and approached expected phenotypic variation in large flocks. This pattern was not as clear for foraging height and foraging behavior. We then examined a dataset of 55 flock matrices from 24 sites across the world. We found that sites with smaller flocks had higher values of phenotypic clumping for body size and sites with larger flocks were less phenotypically clumped. This relationship was weakly negative for foraging behavior and not statistically significant for taxonomic relatedness. Unlike most single-species groups, participants in mixed-species flocks appear to be able to separate on different axes of trait similarity. They can gain benefits from similarity on one axis while mitigating competition by dissimilarity on others. Consistent with our results, we speculate that flock assembly was deterministic up to a certain point with participants being similar in body size, but larger flocks tended to approach random phenotypic assemblages of species.