Mixed-species Flocks Research Articles

On the importance of accounting for alternative foraging tactics when assessing cognitive performance in wild animals.

Research Highlight: Reichert, S., Morand-Ferron, J., Kulahci, I. G., Firth, J. A., Davidson, G. L., Crofts, S. J., & Quinn, J. L. (2021) Cognition and covariance in the producer-scrounger game. Journal of Animal Ecology, https://doi.org/10.1111/1365-2656.13551. When foraging in groups, individuals can either acquire their own resources, as producers, or profit from the work of other individuals, as scroungers. Individuals vary in how much they rely on one foraging tactic over the other. Yet, each of these foraging tactics presents unique cognitive challenges. Using a field experiment with a mixed-species flock of birds, Reichert et al. (2021) investigated how production learning (i.e. successfully feeding from their assigned rewarded feeder) and scrounging propensity (i.e. collecting food from a non-assigned feeders by following another individual) are related at an individual level, as well as the repeatability of both production learning and scrounging propensity. The authors show that overall, (a) individuals learned to scrounge, (b) individuals who rely more on scrounging took longer to learn their assigned feeder and (c) variation in each cognitive trait was mostly explained by individual behavioural flexibility rather than by consistent differences between individuals. Since learning was negatively correlated with the use of an alternative foraging tactic (i.e. scrounging), results of this study also suggest that individual choice of foraging tactics should be considered when evaluating cognitive abilities in wild animals.

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

BackgroundMixed-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.ResultsIn 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).ConclusionsDrongos 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.

Flock-species richness influences node importance and modularity in mixed-species flock networks.

Interdependencies in social groups of animals are a combination of multiple pairwise interactions. Heterospecific groups are often characterized by important species that contribute more to group initiation, maintenance or function than other species. However, in large heterospecific groups, many pairwise interactions are not realised, while others may not be biologically significant, confounding inferences about species importance. Hence, in this study, we examine context dependent changes in species importance and assortment in mixed-species bird flocks from a tropical field site in Southern India using social network analysis. Specifically, we ask how the structural importance of a species and the clustering patterns of species relationships depends on species richness in mixed-species flocks. We constructed both raw and filtered networks; while our results are largely correlated, we believe that filtered networks can provide insights into community-level importance of species in mixed-flocks while raw networks depict flock-level patterns. We find significant differences in flocks of different richness in that different species emerge as structurally important across flocks of varying richness. We also find that assortment is higher in two-species flocks and decreases withan increase in the number of species in the flock ('flock richness' hereafter). We argue that the link between structural importance of species in mixed-species flock networks and their functional significance in the community critically depends on the social context: namely, the species richness of the mixed-species flock. We propose that examining species structural importance at different flock-richness values provides insights into biologically meaningful functional roles of species. More generally, we suggest that it is important to consider context when interpreting species centrality and importance in network structure.

Using playback to test leadership in mixed-species flocks and compare flocking with mobbing

Playback can help identify keystone species in communities that interact behaviourally. Specifically, playback has been used to identify leader species in mixed-species flocks (MSFs) of birds, although it is not clear whether responding heterospecifics are attracted to the leader species or the whole MSF. Playback can also simulate a mobbing response; both flocking and mobbing are regarded as adaptations to predation, but the species participating in these phenomena are not often compared. We performed a complex experiment, comparing heterospecific responses to playback of (1) MSFs (multiple species calling simultaneously), (2) a primary leader species, (3, 4) two secondary leader species that occasionally lead MSFs, (5, 6) two follower species, (7) one nonflocking species and (8) one predator (a small owlet). In 240 trials over 2 years, 72 species made 980 heterospecific approaches to the speaker. We found the predator treatment attracted the most species, and the weakest response was to the nonflocking species. Other patterns were unexpected, however: the response to the MSF, and to the primary leader, was not higher than to other treatments that included MSF members, and the response to one follower species (the particularly vigilant grey-headed canary-flycatcher, Culicicapa ceylonensis) was higher than that to one secondary leader. Although flocking propensity did not influence the response, species that followed MSFs responded more than those that led them. There was a strong correlation between which bird species responded to the owlet and which responded to any MSF member. Our results reveal (1) a response by flock followers to all MSF participants, as if they used these species’ vocalizations to search for the whole MSF, (2) some weak evidence that vigilant species are particularly attractive, and (3) a general correspondence between the species that are attracted to an MSF and those that mob.

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

AbstractWe 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.

Ecology and behavior predict an evolutionary trade‐off between song complexity and elaborate plumages in antwrens (Aves, Thamnophilidae)

The environment can impose constraints on signal transmission properties such that signals should evolve in predictable directions (Sensory Drive Hypothesis). However, behavioral and ecological factors can limit investment in more than one sensory modality leading to a trade-off in use of different signals (Transfer Hypothesis). In birds, there is mixed evidence for both sensory drive and transfer hypothesis. Few studies have tested sensory drive while also evaluating the transfer hypothesis, limiting understanding of the relative roles of these processes in signal evolution. Here, we assessed both hypotheses using acoustic and visual signals in male and female antwrens (Thamnophilidae), a species-rich group that inhabits diverse environments and exhibits behaviors, such as mixed-species flocking, that could limit investment in different signal modalities. We uncovered significant effects of habitat (sensory drive) and mixed-species flocking behavior on both sensory modalities, and we revealed evolutionary trade-offs between song and plumage complexity, consistent with the transfer hypothesis. We also showed sex- and trait-specific responses in visual signals that suggest both natural and social selection play an important role in the evolution of sexual dimorphism. Altogether, these results support the idea that environmental (sensory drive) and behavioral pressures (social selection) shape signal evolution in antwrens.

Structure and Home Range Size of Mixed-Species Bird Flocks in a Bamboo Forest in Southwestern Amazonia

Mixed-species flocks of birds are cohesive groups of two or more different species that are kept in formation by systematic behavioral interactions among the members of the flocks. Typically, mixed-species flocks form themselves around a nuclear species, which presents a specific pattern of movement during foraging, and a song that is recognized by the other species. In the understory of Neotropical forests, representatives of the family Thamnophilidae, in particular species of the genus Thamnomanes, have become adapted for the role of nuclear species in mixed-species flocks. In the present study, we describe the mixed-species flocks led by individuals of Bluish-slate Antshrike Thamnomanes schistogynus in a forest dominated by bamboo Guadua sp. in southwestern Brazilian Amazonia. We collected data within a 600 × 600 m grid subdivided into 36 1-ha plots, with a total area of 36 ha. Within this grid, we delimited the home ranges and core areas of the resident mixed-species flocks. We also identified the composition of each flocks, frequency, fidelity of its members and determined the relationship between mixed-species flocks and vegetation structure within the grid. We delimited the home ranges of eight mixed-species flocks. The mean home range, estimated using the Minimum Convex Polygon method, was 3.45 ± 0.17 ha, and estimated using the autocorrelated kernel density estimation method was 3.65 ± 0.15 ha, and the mean core area, 1.14 ± 0.03 ha. The home ranges of the mixed-species flocks led by T. schistogynus in southwestern Amazonia are among the smallest recorded in species of the genus Thamnomanes. We identified 71 different bird species in the mixed-species flocks with a mean of 41.1 ± 2.1. While the geographic distribution of the lead species T. schistogynus coincides almost exactly with that of the bamboo forests of southwestern Amazonia, our results did not indicate any systematic relationship or fidelity between the mixed-species flocks led by this species and the bamboo forest.

Cognition and covariance in the producer-scrounger game.

The producer-scrounger game is a key element of foraging ecology in many systems. Producing and scrounging typically covary negatively, but partitioning this covariance into contributions of individual plasticity and consistent between individual differences is key to understanding population-level consequences of foraging strategies. Furthermore, little is known about the role cognition plays in the producer-scrounger game. We investigated the role of cognition in these alternative foraging tactics in wild mixed-species flocks of great tits and blue tits, using a production learning task in which we measured individuals' speed of learning to visit the single feeder in an array that would provide them with a food reward. We also quantified the proportion of individuals' feeds that were scrounges ('proportion scrounged'); scrounging was possible if individuals visited immediately after a previous rewarded visitor. Three learning experiments-initial and two reversal learning-enabled us to estimate the repeatability and covariance of each foraging behaviour. First, we examined whether individuals learned to improve their scrounging success (i.e. whether they obtained food by scrounging when there was an opportunity to do so). Second, we quantified the repeatability of proportion scrounged, and asked whether proportion scrounged affected production learning speed among individuals. Third, we used multivariate analyses to partition within- and among-individual components of covariance between proportion scrounged and production learning speed. Individuals improved their scrounging success over time. Birds with a greater proportion scrounged took longer to learn their own rewarding feeder. Although multivariate analyses showed that covariance between proportion scrounged and learning speed was driven primarily by within-individual variation, that is, by behavioural plasticity, among-individual differences also played a role for blue tits. This is the first demonstration of a cognitive trait influencing producing and scrounging in the same wild system, highlighting the importance of cognition in the use of alternative resource acquisition tactics. The results of our covariance analyses suggest the potential for genetic differences in allocation to alternative foraging tactics, which are likely species- and system-dependent. They also point to the need to control for different foraging tactics when studying individual cognition in the wild.

Disturbed Amazonian forests support diminished breeding bird communities

Abstract Twenty percent of the Brazilian Amazon has now been deforested, and deforestation rates are increasing. This compels us to evaluate the conservation potential of human-modified landscapes; yet the ecological value of regenerating and fragmented Amazonian forests remains poorly understood. To date, most faunal studies in disturbed forests have examined metrics derived from presence or abundance. Although valuable, these data cannot tell us how old-growth species are using disturbed forests. In this study, we complement presence data with breeding observations to assess more comprehensively the habitat quality of disturbed forests in central Amazonia. We synthesized breeding evidence from standardized passive mist-netting, mixed-species flock-following, and opportunistic nest-searching across a full annual cycle in 30–35-year-old secondary forest, 10-ha fragments, and undisturbed forest. We then used multistate occupancy models to estimate the number of undetected breeding species in each forest type, which illustrated that fewer species bred in secondary forest (–43%) and 10-ha fragments (–17%) than in undisturbed forest. Both these losses are larger than the associated decrease in species richness alone (–17% and –10%, respectively). Notably, we confirmed breeding by only three terrestrial and near-ground insectivores in fragments and secondary forest combined (of the 9 species found in undisturbed forest). Disturbed forests also supported fewer breeding individuals (–35% to –50%) and, in secondary forest, fewer successful breeding attempts (–24%). Encouragingly, however, some forest-dependent birds are breeding and producing fledglings in disturbed forests, including representatives from almost every guild. This was especially apparent for mixed-species flocks and army-ant followers, two guilds that have historically been considered vulnerable to anthropogenic disturbance. Therefore, despite a loss of breeding habitat in disturbed forests, these data suggest that landscapes composed of regenerating forest and small fragments have conservation potential for forest bird populations.

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

AbstractMixed-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.

Effect of temperature on flocking propensity and species interactions in a subtropical mixed-species flocking system

Increased participation in mixed-species flocks is one hypothesized behavioral tactic used by forest birds to adjust to cold temperatures in winter. This phenomenon has been documented in temperate North America, but not at subtropical latitudes where temperatures are less extreme. To understand how temperature structures the size, richness, flocking propensity, and species interactions of subtropical mixed-species flocks, we performed paired mixed-species flock and point count surveys in upland hardwood forests in north-central Florida, USA. We described the composition of 92 mixed-species flocks across a ∼20 °C range of daytime temperatures, using linear mixed models and network analyses to test for effects of temperature. We found that both flock species richness and size were significantly smaller at higher temperatures while statistically controlling for effects of time of day, canopy height, and local site effects. Thirteen of 14 flocking species showed decreases in within-flock abundance with increasing temperature, although this effect was only significant for the nuclear Carolina Chickadee (Poecile carolinensis). Flock composition and species flocking propensities were less plastic than in temperate systems. By comparing metrics from 7 social networks corresponding to temperature ranges, we found that strength of flocking interactions was lower and that species were significantly less likely to co-occur with similarly foraging species at temperatures above 20 °C. Flocking propensities were lower for many species in the warmest temperature range. Thirteen of 14 species joined flocks in all 7 networks, and dissimilarity of social networks was consequently best explained by changes in species co-occurrence patterns rather than species turnover within flocks. In contrast to boreal flocking systems, changes to flocking behavior were prompted by unseasonably warm weather rather than cold spells, which suggests that warming winter temperatures could lead to breakdown of social behaviors in our system.

Integumentary Colour Allocation in the Stork Family (Ciconiidae) Reveals Short-Range Visual Cues for Species Recognition

The family Ciconiidae comprises 19 extant species which are highly social when nesting and foraging. All species share similar morphotypes, with long necks, a bill, and legs, and are mostly coloured in the achromatic spectrum (white, black, black, and white, or shades of grey). Storks may have, however, brightly coloured integumentary areas in, for instance, the bill, legs, or the eyes. These chromatic patches are small in surface compared with the whole body. We have analyzed the conservatism degree of colouration in 10 body areas along an all-species stork phylogeny derived from BirdTRee using Geiger models. We obtained low conservatism in frontal areas (head and neck), contrasting with a high conservatism in the rest of the body. The frontal areas tend to concentrate the chromatic spectrum whereas the rear areas, much larger in surface, are basically achromatic. These results lead us to suggest that the divergent evolution of the colouration of frontal areas is related to species recognition through visual cue assessment in the short-range, when storks form mixed-species flocks in foraging or resting areas.

The evolution of wood warbler flight calls: Species with similar migrations produce acoustically similar calls.

Diverse animal species engage in long-distance migrations. Many migrants travel in groups, and communication within these groups may be important to survival and successful migration. We examined migration and communication in 36 species of wood warblers (Parulidae), songbirds that breed in North America and migrate in mixed-species flocks to their wintering grounds. During migration, wood warblers produce short vocalizations called "flight calls." The function of flight calls and the patterns of acoustic similarity between species are poorly understood. We investigated whether acoustic similarity of flight calls of different species of warbler reflects the similarity in their migratory journeys or their phylogenetic relatedness. We found that phylogeny, similarity in breeding latitude, and overlap in the timing of migration predict acoustic flight call similarity across warbler species. Further, we found that phylogeny, similarity in migration distance, and overlap in wintering range predict acoustic flight call similarity in a subset of 12 species with highly similar calls, although this analysis has a small sample size. We conclude that migratory similarity may be an important force driving the evolution of acoustically similar calls in wood warblers, in addition to phylogenetic relatedness. Acoustic convergence in these species may facilitate communication between individuals with similar migrations.

Seasonal dynamics of flock interaction networks across a human-modified landscape in lowland Amazonian rain forest.

Although lowland tropical rain forests were once widely believed to be the archetype of stability, seasonal variation exists. In these environments, seasonality is defined by rainfall, leading to a predictable pattern of biotic and abiotic changes. Only the full annual cycle reveals niche breadth, yet most studies of tropical organisms ignore seasonality, thereby underestimating realized conditions. If human-modified habitats display more seasonal stress than intact habitats, then ignoring seasonality will have particularly important repercussions for conservation. We examined the seasonal dynamics of Amazonian mixed-species flocks, an important species interaction network, across three habitats with increasing human disturbance. We quantified seasonal space use, species richness and attendance, and four ecological network metrics for flocks in primary forest, small forest fragments, and regenerating secondary forest in central Amazonia. Our results indicate that, even in intact, lowland rain forest, mixed-species flocks exhibit seasonal differences. During the dry season, flocks included more species, generally ranged over larger areas, and displayed network structures that were less complex and less cohesive. We speculate that-because most flocking species nest during the dry season, a time of reduced arthropod abundance-flocks are simultaneously constrained by these two competing pressures. Moreover, these seasonal differences were most pronounced in forest fragments and secondary forest, habitats that are less buffered from the changing seasons. Our results suggest that seasonality influences the conservation value of human-modified habitats, raising important questions about how rain forest organisms will cope with an increasingly unstable climate.

Is Intestinal Bacterial Diversity Enhanced by Trans-Species Spread in the Mixed-Species Flock of Hooded Crane (Grus monacha) and Bean Goose (Anser fabalis) Wintering in the Lower and Middle Yangtze River Floodplain?

Simple SummaryIntestinal microbes play an indispensable role in host physiology and their alteration can produce serious effects on vertebrates. In this study, we analyzed the characteristics of intestinal bacterial community of hooded crane and bean goose whose niches overlap at Shengjin Lake, China, and investigated how host internal factors and inter-species interactions affected the diversity and spread of intestinal bacteria of the two species over three wintering periods. We have found that direct or indirect contact with each other increased the diversity of host intestinal bacteria and caused bacteria to spread among species in the mixed-species flock. In addition, a total of 63 pathogens were identified, of which 38 (60.3%) were found in the gut of both species. These findings could help our understanding of the factors that influence gut bacteria in wild waterbirds, which are also major contributors to the spread of pathogens worldwide.Diversity of gut microbes is influenced by many aspects, including the host internal factors and even direct or indirect contact with other birds, which is particularly important for mixed-species wintering waterbird flocks. In this study, Illumina high-throughput sequencing was used to analyze the intestinal bacteria of the hooded crane and bean goose whose niches overlap at Shengjin Lake. We tested whether contact time enhances the trans-species spread of gut bacteria. Results indicate alpha-diversity and microbial composition displayed significant separation between the two hosts in every wintering period, although the number of bacteria types shared increased with increasing contact time. For the same species, with the lengthening of contact time, alpha-diversity and the number of operational taxonomic units (OTUs) in the host intestine augmented, and the common OTUs and structural similarity of microflora in the middle and late periods were more than in the early and middle periods. In addition, we found a very high proportion of shared pathogens. Our results indicate that, although intestinal microflora of different species were separated, direct or indirect contact in the mixed-species flock caused the spread of gut bacteria trans-species, indicating that more attention should be paid to intestinal pathogens in wild birds.

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.

Neighborhood effects in mixed-species flocks affect foraging efficiency of intermediate and little egrets

Birds aggregate for various survival needs, and ardeids form both conspecific and mixed-species flocks. However, it remains unclear whether and how ardeids’ foraging while joining a flock is affected differently in conspecific versus mixed-species flocks. We studied egret flocks in southwestern Taiwan to assess the flock size and neighborhood effects on flock formation and foraging performance of egrets by examining landing tendencies and expelling behaviors of Little (Egretta garzetta) and Intermediate (E. intermedia) egrets, and assessing strike rates, strike efficiency, and intake rates of singles and egrets in conspecific and mixed-species flocks. We found no evidence of non-random landings to mixed-species flocks by either species, but egrets showed a trend of landing closer to conspecifics when conspecifics represented lower proportions in flocks. Intermediate Egrets displayed expelling behavior more frequently. Expelling by earlier arrived egrets interrupted subsequent flock formation on 45.2 % occasions, but was not affected by egrets’ foraging performance. Solitary Little Egrets had higher strike and fish intake rates, but lower strike efficiency, than solitary Intermediate Egrets, whereas their biomass intake rates were similar. Strike and fish intake rates increased in flocks compared to singles in both species, but declined for Little Egrets as flocks increased from medium (11–30 birds) to large sizes (> 30). Intermediate Egrets had higher strike, fish intake, and biomass intake rates, but Little Egrets had lowered strike efficiency and fish- and biomass-intake rates, when near each other than when near conspecifics. Overall, Intermediate Egrets in mixed-species flocks improved foraging efficiency and food intakes by neighborhood effects at costs of reduced foraging efficiency to Little Egrets. The variation in landing tendency over flock compositions suggests potential interactions of the neighborhood effect with egrets’ conspecific attraction.

Vegetation structure drives mixed-species flock interaction strength and nuclear species roles

Abstract Mixed-species flocks are a key facilitative interaction for tropical birds. Forest fragmentation leads to species loss and spatial turnover in these flocks, yet it is unknown how these changes to composition influence within-flock species interactions. We used network analysis to characterize flocking interactions along a fragment-size gradient in the Colombian Western Andes. We asked 1) how patch size, edge density, and vegetation structure explained network measures indicative of flock cohesion, 2) whether changes were driven by flocking species turnover or changes to the frequency of species co-occurrence, and 3) whether nuclear species, those that maintain flock stability and cohesion, changed in importance across the gradient. We constructed weighted social networks from flock compositions observed on 500-m transects, and then calculated global network measures and the centrality of six nuclear species. Patch size and edge density did not correlate with interspecific co-occurrence patterns, but interaction strength increased with canopy height. Flocks contained numerous, weak interactions, and there were no flock subtypes, suggesting flock composition was dynamic and unstructured. Several redundant nuclear species were present and varied in importance based on ecological conditions. A chlorospingus (Passerellidae) was most central in old-growth forest, whereas several tanager (Thraupidae) species became more central in smaller fragments and disturbed forest. When partitioning network dissimilarity, we found that 66% of dissimilarity resulted from species turnover, whereas only 34% resulted from changes to species co-occurrence. This finding suggests that coherence of flocking behavior itself is maintained even as extensive species turnover occurs from continuous forest to small fragments.

Do racket-tailed drongos make tree guards for their nest trees?

Abstract Greater racket-tailed drongos are renowned for their splendid mimicking abilities, and for their significant roles within mixed species flocks in the Old World tropics. Yet, we know little about their basic ecology and breeding behaviour. Here we describe a set of unique behaviours of these drongos during their nesting season. Racket-tailed drongos nested in trees in an open patch of forest, often returning to the same tree year after year. The nesting pair also smoothened the bark of the nest tree trunk with their beaks. These findings suggest that the nest tree is a crucial resource for this species, and have implications for the cognitive abilities of drongos, as well as for hitherto unknown interactions between an avian species and tropical forest trees.

The relationship between morphology and behavior in mixed-species flocks of island birds.

Understanding how co‐occurring species divide ecological space is a central issue in ecology. Functional traits have the potential to serve as a means for quantitatively assessing niche partitioning by different species based on their ecological attributes, such as morphology, behavior, or trophic habit. This enables testing ecological and evolutionary questions using functional traits at spatio‐temporal scales that are not feasible using traditional field methods. Both rapid evolutionary change and inter‐ and intraspecific competition, however, may limit the utility of morphological functional traits as indicators of how niches are partitioned. To address how behavior and morphology interact, we quantified foraging behavior of mixed‐species flocks of birds in the Solomon Islands to test whether behavior and morphology are correlated in these flocks. We find that foraging behavior is significantly correlated with morphological traits (p = .05), but this correlation breaks down after correcting for phylogenetic relatedness (p = .66). These results suggest that there are consistent correlations between aspects of behavior and morphology at large taxonomic scales (e.g., across genera), but the relationship between behavior and morphology depends largely on among‐clade differences and may be idiosyncratic at shallower scales (e.g., within genera). As a result, general relationships between behaviors and morphology may not be applicable when comparing close relatives.