Mixed-species Flocks Research Articles

Mixed-species flock sizes and compositions influence flock members' success in three field experiments with novel feeders.

Mixed-species groups and aggregations are quite common and may provide substantial fitness-related benefits to group members. Individuals may benefit from the overall size of the mixed-species group or from the diversity of species present, or both. Here we exposed mixed-species flocks of songbirds (Carolina chickadees, Poecile carolinensis, tufted titmice, Baeolophus bicolor, and the satellite species attracted to these two species) to three different novel feeder experiments to assess the influence of mixed-species flock size and composition on ability to solve the feeder tasks. We also assessed the potential role of habitat density and traffic noise on birds' ability to solve these tasks. We found that likelihood of solving a novel feeder task was associated with mixed-species flock size and composition, though the specific social factor involved depended on the particular species and on the novel feeder. We did not find an influence of habitat density or background traffic noise on likelihood of solving novel feeder tasks. Overall, our results reveal the importance of variation in mixed-species group size and diversity on foraging success in these songbirds.

Dietary DNA metabarcoding reveals a trophic niche partitioning among sympatric Iberian sandgrouses and bustards

The study of trophic niche partitioning is of great importance for understanding community structure and species coexistence, particularly if these are threatened. Here DNA metabarcoding was used to assess the diet of four threatened steppe bird species (two bustards and two sandgrouses), with the aim of better understanding their dietary requirements, trophic interactions, and potential threats. The results showed seasonal and interspecific differences in their plant diet, with greater importance of cultivated plants during autumn and winter (around 50 % of their diet) than spring. Plants of the genus Convolvulus and of the family Brassicaceae were frequently consumed by all species. In spring, poppies (Papaver spp.) were a considerable part of their diet, and could be used as a source of carotenoids or for their anti-parasitic properties. Furthermore, results evidenced a trophic niche partitioning among species, with a marked segregation between bustard species and, to a lesser extent, between sandgrouse species. Diet similarity was generally higher between species from different orders that occur in mixed-species flocks (bustard - sandgrouse) than between species of the same order. This partitioning was probably related to a stratification in habitat use rather than to specialisation and might prevent competition to some extent. However, the homogenization of trophic resources resulting from agricultural intensification could pose an important threat, particularly during autumn, when weeds are scarcer and the most abundant trophic resource are sown seeds, which are often treated with pesticides.

Strangers like me: birds respond equally to a familiar and an unfamiliar sentinel species' alarm calls, but respond less to non‐core and non‐sentinel's alarm calls

Alarm signals have evolved to communicate imminent threats to conspecifics but animals may also perceive other species' alarm displays to obtain adaptive information. In birds, mixed‐species foraging flocks are often structured around a focal sentinel species, which produces reliable alarm calls that inform eavesdropping non‐sentinel heterospecifics about predation risk. Ongoing work has revealed that several species can recognize the alarm calls of certain sentinel species even without prior encounters, including when these are from distant biogeographic regions. Similar work has yet to examine whether naive subjects' responses to unfamiliar sentinel alarm calls differ from responses to non‐sentinel alarm calls. Here we played the alarm calls of three subtropical Asian bird species that participate in mixed species flocks, to temperate North American birds. Birds responded most to the alarm call of an allopatric core sentinel and a local sympatric sentinel control species, less so to an allopatric non‐core sentinel, and least so to an allopatric non‐sentinel and a negative control stimulus. These patterns provide evidence that broad phylogenetic and geographic recognition is a pertinent aspect of sentinel alarm calls in general.

Variations in inter-specific and sex-related niche partitioning in pelagic boobies during their annual cycle.

Animals that co-occur in a region (sympatry) may share the same environment (syntopy), and niche differentiation is expected among closely related species competing for resources. The masked booby (Sula dactylatra) and smaller congeneric red-footed booby (Sula sula) share breeding grounds. In addition to the inter-specific size difference, females of both species are also larger than the respective males (reversed sexual size dimorphism). Although both boobies consume similar prey, sometimes in mixed-species flocks, each species and sex may specialize in terms of their diet or foraging habitats. We examined inter- and intra-specific differences in isotopic values (δ13C and δ15N) in these pelagically feeding booby species during the incubation period at Clarion Island, Mexico, to quantify the degrees of inter- and intra-specific niche partitioning throughout the annual cycle. During incubation, both species preyed mainly on flyingfish and squid, but masked boobies had heavier food loads than red-footed boobies. There was no overlap in isotopic niches between masked and red-footed boobies during breeding (determined from whole blood), but there was slight overlap during the non-breeding period (determined from body feathers). Female masked boobies had a higher trophic position than conspecific males during breeding; however, no such pattern was detected in red-footed boobies. These results provide evidence of inter- and intra-specific niche partitioning in these tropical seabird species, particularly during the breeding period and in the more-dimorphic species. Our results suggest that these closely related species use different strategies to cope with the same tropical marine environment.

Investigating flock-associated mimicry: examining the evidence for, and drivers of, plumage mimicry in the greater and lesser necklaced laughingthrush.

Visual mimicry is less understood in birds than in other taxa. The interspecific social dominance mimicry (ISDM) hypothesis asserts that subordinate species resemble dominant ones to reduce aggression. Plumage mimicry has also been consistently noted in mixed-species flocks (MSFs), suggesting a connection to grouping behaviour, although it is unclear whether this is linked to ISDM. We studied greater necklaced laughingthrush (GNLT, Pterorhinus pectoralis) and lesser necklaced laughingthrush (LNLT, Garrulax monileger), which were recently placed in different genera. Measurements of 162 museum specimens showed LNLT converging in sympatry with GNLT in necklace colour, but diverging in necklace to body ratio, with proportionally smaller necklaces. The species were closely associated in six of seven MSF systems from Nepal to China. In a study of foraging behaviour in Nepal, aggression was rare between the species, LNLT followed GNLT and had lower foraging rates when further from GNLT. Our data suggest a link between this MSF-associated mimicry and ISDM, and that the subordinate LNLT may be the mimic and gain more from the resemblance. The species spend much time together in dense and poorly lit vegetation, where the LNLTs resemblance to GNLTs potentially allows them to forage closer to GNLTs than would be otherwise possible.

Sharp declines in observation and capture rates of Amazon birds in absence of human disturbance

Across the globe, unprecedented declines have been reported across a range of taxa. Among the most well documented are declines in bird populations, with most declines attributable to human activities such as deforestation and other alterations to habitats. There is increasing evidence that bird populations also have declined at sites within large expanses of relatively undisturbed lowland tropical forest, such as in Amazonia. Causes of such declines likely are varied and may be related to direct or indirect effects of climate change. Here, we use mist-net data and direct observations to examine changes in bird numbers over a 22-yr period on two 100-ha study plots in an Amazonian lowland forest of eastern Ecuador. As previously reported, capture rates experienced large declines from 2009 on; observation rates also showed large declines through 2013 but subsequently have shown no consistent trend up or down. Overall capture rates and observation rates are now approximately half the rates during the first decade. Most foraging guilds also declined over time, with declines particularly pronounced among insectivorous groups. With few exceptions, patterns of change were similar between plots suggesting a more general response rather than responses to local factors on each plot. Capture rates and observation rates of ∼90% of the most common species in both understory and canopy were lower during the latter part of the study, with only a few species (∼10%) showing increases. Similarly, large declines were seen in both canopy and understory mixed-species flocks. Large declines in bird numbers have been noted in other tropical sites that have conducted long-term population studies, including Panama and Brazil, suggesting a response to large-scale factors, such as climate change. While not yet documented, the consequences of losing approximately 50% of population abundance are likely substantial and could result in local extinction of rare species, altered species interactions and social organization, as well as declining ecological function and biotic integrity of these tropical forest ecosystems.

Convergent niche shifts of endangered parrots (genus Amazona) during successful establishment in urban southern California

AbstractAimIntroduced species offer insight on whether and how organisms can shift their ecological niches during translocation. The genus Amazona offers a clear test case, where sister species Red‐crowned (A. viridigenalis) and Lilac‐crowned Parrots (A. finschi) have established breeding populations in southern California following introduction via the pet trade from Mexico where they do not coexist. After establishment in the 1980s, introduced population sizes have increased, with mixed species flocks found throughout urban Los Angeles. Here, we investigate the differences between the environmental conditions of the native and introduced ranges of these now co‐occurring species.LocationSouthern California and Mexico.MethodsUsing environmental data on climate and habitat from their native and introduced ranges, we tested whether Red‐crowned and Lilac‐crowned Parrots have divergent realized niches between their native ranges, and whether each species has significantly shifted its realized niche to inhabit urban southern California. We also analysed data from Texas and Florida introductions of Red‐crowned Parrots for comparative analysis.ResultsThere are significant differences in the native‐range niches of both parrot species, but a convergence into a novel, shared environmental niche into urban southern California, characterized by colder temperatures, less tree cover and lower rainfall. Texas and Florida Red‐crowned Parrots also show evidence for niche shifts with varying levels of niche conservatism through the establishment of somewhat different realized niches.Main ConclusionsDespite significant niche shifts, introduced parrots are thriving, suggesting a broad fundamental niche and an ability to exploit urban resources. Unique niche shifts in different U.S. introductions indicate that Amazona parrots can adapt to diverse environmental conditions, with cities offering a resource niche and the timing of introduction playing a crucial role. Cities can potentially serve as refugia for threatened parrot species, but the risk of hybridization between species emphasizes the need for ongoing monitoring and genetic investigations.

The impacts of host traits on parasite infection of montane birds in southwestern China

Parasitic infections have the potential to impact the hosts’ body condition, elevate physiological responses, and ultimately lead to increased mortality. Host-parasite interactions are tied to the ecological and life-history traits of the hosts. While montane birds are susceptible to avian blood parasites, few studies have simultaneously assessed how inter- and intra-specific traits of hosts influence their probability of parasite infection. In this study, we screened for avian blood parasites across 214 individuals from 51 species at two sites, including a lowland farmland at 700 m and a highland forest at 2,500 m, in the Gaoligong Mountains in southwestern China. Overall blood parasite prevalence was 53.74%, with divergent species-specific prevalence ranging from 6.25% to 66.67%. We also measured traits indicative of body condition and physiological responses of each sampled individual. Using Bayesian phylogenetic logistic models, we assessed whether parasite infection probability is associated with ecological and life history traits of host species. Larger bird species were more likely to be infected than smaller bird species, and omnivore species showed lower susceptibility than those with other diets such as insectivores and herbivores. In contrast, foraging strata, nest type, and participation in mixed-species flocks of host species did not affect infection probability. We then used a reduced sample of eight species with more than five individuals, to assess the associations between intra-specific infection probability and host body condition, represented by fat and muscle reserves, and acute stress responses measured through breath rate. While infected individuals were likely to have more fat reserves than non-infected individuals, we did not find any association between infection probability and muscle reserve and body mass, nor with breath rate. Our results revealed that at the species level, specific traits (body mass and diet) of host species predict infection probability and implied a potential link at the individual level between enhanced body condition and increased resilience to parasite infection.

Influence of social and physical environmental variation on antipredator behavior in mixed-species parid flocks.

Carolina chickadees (Poecile carolinensis) and tufted titmice (Baeolophus bicolor) regularly form flocks with multiple species through the winter months, including white-breasted nuthatches (Sitta carolinensis). Earlier studies found that behavior of both chickadees and titmice was sensitive to mixed-species flock composition. Little is known about the influence of background noise level and vegetation density on the antipredator behaviors of individuals within these flocks, however. We tested for the effects of vegetation density, traffic noise, and flock composition (conspecific number, flock diversity, and flock size) on antipredator behavioral responses following an alarm call playback (Study 1) and an owl model presentation (Study 2) at feeders. We recorded background traffic noise and performed lidar scans to quantify vegetation density at each site. After a feeder had been stocked with seed and a flock was present, we recorded calls produced, and we identified flock composition metrics. We coded seed-taking latency, call latency, mob latency, and mob duration following the respective stimulus presentation and tested for effects of flock composition metrics, vegetation density, and background noise on these responses. For the alarm call playback study, flock composition drove behaviors in chickadees and titmice, and vegetation density drove behaviors in chickadees and nuthatches. For the owl model study, conspecific number predicted behavior in chickadees, and mob duration was predicted by nuthatch number. The results reveal individual sensitivity to group composition in anti-predatory and foraging behavior in simulated risky contexts. Additionally, our data suggest that the modality of perceived simulated risk (acoustic vs. visual) and the density of vegetation influence behavior in these groups.

Human-facilitated feeding on Date Palm Phoenix dactylifera sap by trans-Saharan warblers during spring stopover in a Tunisian oasis

ABSTRACT Capsule Some trans-Saharan warbler species feed on human-extracted Date Palm Phoenix dactylifera sap during spring migratory stopover. Aims To investigate the behaviour of birds feeding on Date Palm sap at a Tunisian oasis during the spring migration season. Methods Birds feeding on Date Palm sap were monitored during the peak of spring migratory passage of trans-Saharan birds in southern Tunisia by recording the species involved and their numbers, and noting whether there were agonistic interactions between them. Data were used to describe the assemblages of birds feeding on Date Palm sap. Results We recorded five trans-Saharan warbler species feeding on Date Palm sap, either in single- or mixed-species flocks. Aggressive conflicts were common, suggesting competition by interference within this bird assemblage. However, other abundant fruit-eating trans-Saharan warbler species were never seen feeding on Date Palm sap, possibly due to the potential inability to assimilate some of its components, or simply because they did not need to ingest it for reasons related to their migration strategy. Conclusions Our findings reveal that Date Palm sap is a food resource for some trans-Saharan warbler species, highlighting a rarely documented human-facilitated feeding behaviour in birds. They also underline the high ecological plasticity of these warbler species, capable of using different food resources to meet the needs of their challenging migratory journey.

What's the rumpus? Resident temperate forest birds approach an unfamiliar neotropical alarm call across three continents.

Alarm signals have evolved to communicate pertinent threats to conspecifics, but heterospecifics may also use alarm calls to obtain social information. In birds, mixed-species flocks are often structured around focal sentinel species, which produce reliable alarm calls that inform eavesdropping heterospecifics about predation risk. Prior research has shown that Neotropical species innately recognize the alarm calls of a Nearctic sentinel species, but it remains unclear how generalizable or consistent such innate signal recognition of alarm-calling species is. We tested for the responses to the alarm calls of a Neotropical sentinel forest bird species, the dusky-throated antshrike (Thamnomanes ardesiacus), by naive resident temperate forest birds across three continents during the winter season. At all three sites, we found that approaches to the Neotropical antshrike alarm calls were similarly frequent to the alarm calls of a local parid sentinel species (positive control), while approaches to the antshrike's songs and to non-threatening columbid calls (negative controls) occurred significantly less often. Although we only tested one sentinel species, our findings indicate that temperate forest birds can recognize and adaptively respond globally to a foreign and unfamiliar tropical alarm call, and suggest that some avian alarm calls transcend phylogenetic histories and individual ecological experiences.

Questionnaire study suggests grave consequences of infectious laryngotracheitis, infectious coryza and mycoplasmosis in small chicken flocks

BackgroundA growing number of people in western countries keep small chicken flocks. In Sweden, respiratory disease is a common necropsy finding in chickens from such flocks. A respiratory real-time polymerase chain reaction (PCR) panel was applied to detect infectious laryngotracheitis virus (ILTV), Avibacterium paragallinarum (A. paragallinarum) and Mycoplasma gallisepticum (M. gallisepticum) in chickens from small flocks which underwent necropsy in 2017–2019 and had respiratory lesions. Owners (N = 100) of PCR-positive flocks were invited to reply to a web-based questionnaire about husbandry, outbreak characteristics and management.ResultsResponse rate was 61.0%. The flocks were from 18 out of Sweden’s 21 counties indicating that respiratory infections in small chicken flocks are geographically widespread in Sweden. Among participating flocks, 77.0% were coinfected by 2–3 pathogens; 91.8% tested positive for A. paragallinarum, 57.4% for M. gallisepticum and 50.8% for ILTV. Larger flock size and mixed-species flock structure were associated with PCR detection of M. gallisepticum (P = 0.00 and P = 0.02, respectively). Up to 50% mortality was reported by 63.9% of respondents. Euthanasia of some chickens was carried out in 86.9% of the flocks as a result of the outbreaks. Full clinical recovery was reported by 39.3% of owners suggesting chronic infection is a major challenge in infected flocks. Live birds had been introduced in many flocks prior to outbreaks, which suggested these as an important source of infection. Following the outbreaks, 36.1% replaced their flocks with new birds and 9.8% ceased keeping chickens.ConclusionsThis study highlights the severity of respiratory outbreaks in small non-commercial chicken flocks and points to the need for more research and veterinary assistance to prevent and manage respiratory infections in small chicken flocks.

A classification scheme for mixed-species bird flocks.

The literature on mixed-species flocks references a wide variety of bird associations. These studies, however, have used an array of unstructured characteristics to describe flocks, ranging from the temporal occurrence of flocking to the identity and behavioural features of constituent members, with little consensus on which key traits define and characterize a mixed-species flock. Moreover, although most studies report species-specific roles, there is no clear consensus about what these roles signify nor how to define them. This lack of consistency limits our ability to compare flocks from different habitats, regions and species pools. To unify this sizable body of literature, we reviewed and synthesized 538 studies on mixed-species flocks. We propose 13 categories to classify mixed-species flocks using behavioural and physical traits at the flock and participant level, as well as the habitat where the flock occurs. Lastly, we discuss the historical terminology for different species roles and propose definitions to clarify and distinguish among nuclear, leader, sentinel, and flock-following species. We envision that these guidelines will provide a universal language for mixed-species flock research, paving the way for future comparisons and new insight between different regions and systems. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

Acoustic similarity of flight calls corresponds with the composition and structure of mixed-species flocks of migrating birds: evidence from a three-dimensional microphone array.

Animals that migrate in mixed-species groups may communicate with both conspecific and heterospecific individuals, providing a low-cost mechanism for navigation whenever individuals share similar migratory routes or destinations. Many migratory birds produce calls while flying, but the function of these calls, and the forces contributing to their evolution, are poorly known. We studied flight calls in mixed-species groups of wood warblers (Parulidae), a biodiverse group of migratory songbirds. We used a spatial approach to examine whether acoustic similarity of flight calls varies with group composition, recording flight calls of mixed-species flocks with a wireless microphone array and triangulating the positions of birds in three dimensions. We found that the acoustic similarity of flight calls was correlated with spatial proximity: birds with similar calls fly closer together during migration. We also found relationships between acoustic similarity, flock size and mixed-species flock diversity: birds with similar calls fly in smaller flocks and in flocks with lower species diversity. Our results support the idea that migrating birds use flight calls to maintain contact with acoustically similar individuals in mixed-species flocks, with communication transcending species boundaries. These results suggest that acoustically similar flight calls are used as cues of group assembly for migratory animals. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

Revisiting the 'nuclear species' concept: do we really know what we think we know?

The idea of 'nuclear species' has received a lot of attention in mixed-species flock research. Our impression of this literature is that referenced statements tend to cite the same papers in support of a small set of ideas, and often there is a mismatch between what papers contain and what they're cited for. Motivated by these impressions, we built and quantitatively examined a database of referenced statements about nuclearity in flocks. This confirmed our impression quantitatively, but more strikingly, a single paper stood out in its influence on ideas around nuclearity in flocks. Moynihan's 1962 monograph on mixed-species flocks in Panama, 'The organization and probable evolution of some mixed-species flocks of neotropical birds' published in Smithsonian Miscellaneous Collections, was cited twice as much as the next most-cited paper and was the most-cited paper for 10 out of 15 most-discussed ideas related to nuclearity. Further, a number of other highly cited papers are strongly influenced by Moynihan's ideas, i.e. its influence is much greater than what a count of citations conveys. We also found that Moynihan was mis-cited frequently. We juxtapose what we found from the citation analysis with what the paper actually contains to better understand the nature of support that Moynihan provides, and discuss the implications of our findings for what we know about and how we research nuclearity in flocks. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

Relationship among vegetation structure and mixed-species flocks composition along the latitudinal gradient of the subtropical montane forest of the Yungas, Argentina.

The structural and physiognomic characteristics of forests and mountain forest are fundamental aspects that influence the richness, abundance and composition in the bird community. The objective of the present work was to analyse how the vegetation structure could influence the species composition of mixed-species flocks (MSF), along the latitudinal gradient of the subtropical montane forest of the Yungas of Argentina. Eight sites were studied along the 700 km of distribution of the Argentine Yungas. Richness and abundance of MSF were determined. In addition, different variables of composition and structure of the vegetation were analysed. Multivariate analysis indicated that vertical strata coverage and litter depth were the main variables associated with changes in the species composition of MSF along the gradient. Variation in MSF composition within the Yungas was associated with the physiognomy of the subtropical montane forest, which could indicate that it is strongly linked to the condition of the local vegetation. Substantial changes in vegetation could drastically change the composition of the resident flocks. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

Interspecific sociality alters the colonization and extinction rates of birds on subtropical reservoir islands.

Island biogeography theory has proved a robust approach to predicting island biodiversity on the assumption of species equivalency. However, species differ in their grouping behaviour and are entangled by complex interactions in island communities, such as competition and mutualism. We here investigated whether intra- and/or interspecific sociality may influence biogeographic patterns, by affecting movement between islands or persistence on them. We classified bird species in a subtropical reservoir island system into subcommunities based on their propensity to join monospecific and mixed-species flocks. We found that subcommunities which had high propensity to flock interspecifically had higher colonization rates and lower extinction rates over a 10-year period. Intraspecific sociality increased colonization in the same analysis. A phylogenetically corrected analysis confirmed the importance of interspecific sociality, but not intraspecific sociality. Group-living could enable higher risk crossings, with greater vigilance also linked to higher foraging efficiency, enabling colonization or long-term persistence on islands. Further, if group members are other species, competition can be minimized. Future studies should investigate different kinds of island systems, considering positive species interactions driven by social behaviour as potential drivers of community assembly on islands. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

Can convergence in mixed-species flocks lead to evolutionary divergence? Evidence for and methods to test this hypothesis.

One of the most fundamental goals of modern biology is to achieve a deep understanding of the origin and maintenance of biodiversity. It has been observed that in some mixed-species animal societies, there appears to be a drive towards some degree of phenotypic trait matching, such as similar coloration or patterning. Here we build on these observations and hypothesize that selection in mixed-species animal societies, such as mixed-species bird flocks, may drive diversification, potentially leading to speciation. We review evidence for possible convergent evolution and even outright mimicry in flocks from southwestern China, where we have observed several cases in which species and subspecies differ from their closest relatives in traits that match particular flock types. However, understanding whether this is phenotypic matching driven by convergence, and whether this divergence has promoted biodiversity, requires testing multiple facets of this hypothesis. We propose a series of steps that can be used to tease apart alternative hypotheses to build our understanding of the potential role of convergence in diversification in participants of mixed-species societies. Even if our social convergence/divergence hypothesis is not supported, the testing at each step should help highlight alternative processes that may affect mixed-species flocks, trait evolution and possible convergence. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

Neotropical mixed-species bird flocks in a community context.

Mixed-species flocks are an important component of bird communities, particularly in the Neotropics, where flocks reach their highest diversity. The extent to which mixed-species flocks represent unique functional or ecological roles within communities, and how these attributes change over environmental gradients, however, is not well understood. We use a trait-based approach to examine functional aspects of flocking assemblages as they relate to those observed in the larger avian community across a 3000 m elevational gradient. Our results reveal similar ecological strategies among flocking species and the communities in which they occur, at the scale of the regional pool and across elevations. Trait variation in flocking and non-flocking assemblages is structured along two major axes defined by size- and resource-related traits. The trait space occupied by flocking species, however, represents only half (51%) that of the larger community. Similarly, the trait space of flocks across elevations is restricted compared to non-flocking species. The shared trait space across flock types represents small-bodied invertivores foraging in lower forest strata, traits associated with increased vulnerability to predation. The concentration of flocking species in functional trait space suggests high niche packing and either more overlap in ecological strategies or more finely divided niches relative to non-flocking species. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

Network structure of avian mixed-species flocks decays with elevation and latitude across the Andes

Birds in mixed-species flocks benefit from greater foraging efficiency and reduced predation, but also face costs related to competition and activity matching. Because this cost-benefit trade-off is context-dependent (e.g. abiotic conditions and habitat quality), the structure of flocks is expected to vary along elevational, latitudinal and disturbance gradients. Specifically, we predicted that the connectivity and cohesion of flocking networks would (i) decline towards tropical latitudes and lower elevations, where competition and activity matching costs are higher, and (ii) increase with lower forest cover and greater human disturbance. We analysed the structure of 84 flock networks across the Andes and assessed the effect of elevation, latitude, forest cover and human disturbance on network characteristics. We found that Andean flocks are overall open-membership systems (unstructured), though the extent of network structure varied across gradients. Elevation was the main predictor of structure, with more connected and less modular flocks upslope. As expected, flocks in areas with higher forest cover were less cohesive, with better defined flock subtypes. Flocks also varied across latitude and disturbance gradients as predicted, but effect sizes were small. Our findings indicate that the unstructured nature of Andean flocks might arise as a strategy to cope with harsh environmental conditions. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.