Baeolophus Bicolor Research Articles

Comparative analysis of meningeal transcriptomes in birds: Potential pathways of resilience to repeated impacts.

The meninges and associated vasculature (MAV) play a crucial role in maintaining cerebral integrity and homeostasis. Recent advances in transcriptomic analysis have illuminated the significance of the MAV in understanding the complex physiological interactions at the interface between the skull and the brain after exposure to mechanical stress. To investigate how physiological responses may confer resilience against repetitive mechanical stress, we performed the first transcriptomic analysis of avian MAV tissues using the Downy Woodpecker (Dryobates pubescens) and Tufted Titmouse (Baeolophus bicolor) as the comparison species. Our findings reveal divergences in gene expression profiles related to immune response, cellular stress management, and protein translation machinery. The male woodpeckers exhibit a tailored immune modulation strategy that potentially dampens neuroinflammation while preserving protective immunity. Overrepresented genes involved in cellular stress responses suggest enhanced mechanisms for mitigating damage and promoting repair. Additionally, the enrichment of translation-associated pathways hints at increased capacity for protein turnover and cellular remodeling vital for recovery. Our study not only fills a critical gap in avian neurobiology but also lays the groundwork for research in comparative neuroprotection.

Do vocal mimics learn their mimetic songs from heterospecifics or conspecifics?

Abstract Vocal mimicry can develop either through imitation of heterospecific models (primary mimicry) or through imitation of conspecifics already producing mimetic song (secondary mimicry). Distinguishing primary versus secondary mimicry is important from ecological, evolutionary, and neurobiological perspectives. We outline four empirical strategies for researchers to detect the usage of secondary mimicry: (1) model selection strategy (heterospecific model commonly mimicked but rare or absent locally), (2) usage frequency strategy (syllable types commonly mimicked but rarely used by the model), (3) acoustic structure strategy (acoustic divergence in how a syllable type gets produced by mimics versus models), and (4) syntax strategy (consistent variation between mimic and model in the ordering of syllable types). We then use these strategies to produce evidence for secondary mimicry in northern mockingbirds (Mimus polyglottos), focusing mostly on the mimicked songs of northern cardinals (Cardinalis cardinalis) and tufted titmice (Baeolophus bicolor). As further evidence of the mechanisms by which secondary mimicry might occur, we also demonstrate that mockingbirds match countersing much more frequently in response to conspecifics versus heterospecifics. Deeper questions about the precise extent of primary versus secondary mimicry in mockingbirds and other mimicking species still need answers.

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.

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.

Interspecific differences in the effects of masking and distraction on anti-predator behavior in suburban anthropogenic noise.

Predation is a common threat to animal survival. The detection of predators or anti-predator communication signals can be disrupted by anthropogenic noise; however, the mechanism by which responses are affected is unclear. Masking and distraction are the two hypotheses that have emerged as likely explanations for changes in behavior in noise. Masking occurs when the signal and noise fall within the same sensory domain; noise overlapping the energy in the signal reduces signal detection. Distraction can occur when noise in any sensory domain contributes to a greater cognitive load, thereby reducing signal detection. Here, we used a repeated measures field experiment to determine the relative contributions of masking and distraction in mediating reduced anti-predator responses in noise. We recorded the approaches and vocalizations of black-capped chickadees (Poecile atricapillus), tufted titmice (Baeolophus bicolor), and white-breasted nuthatches (Sitta carolinensis) to both visual and acoustic cues of predator presence, either with or without simultaneous exposure to anthropogenic noise. Titmice increased their calling to both visual and acoustic cues of predator presence. However, there was no significant effect of noise on the calling responses of titmice regardless of stimulus modality. Noise appeared to produce a distraction effect in chickadees; however, this effect was small, suggesting that chickadees may be relatively unaffected by low levels of anthropogenic noise in suburban environments. White-breasted nuthatch calling behavior was affected by the interaction of the modality of the predator stimulus and the noise condition. Nuthatches had a delayed response to the predator presentations, with a greater calling rate following the presentation of the acoustic stimulus in quiet compared to the presentation of the acoustic stimulus in noise. However, there was no difference in calling rate between the quiet and noise conditions for the visual stimulus. Together this suggests that even moderate levels of noise have some masking effect for white-breasted nuthatches. We suggest that the mechanisms through which noise influences anti-predator behavior may depend on the social roles, foraging ecology and auditory capabilities of each species.

Low Incidence of Avian Predation on the Brown Marmorated Stink Bug, Halyomorpha halys (Hemiptera: Pentatomidae), in Southeastern Orchard Systems.

In many agroecosystems, brown marmorated stink bugs (Halyomorpha halys) (Hemiptera: Pentatomidae) are polyphagous pests that cause significant economic losses to numerous crops every year. Insectivorous birds may provide a means of sustainable predation of invasive pests, such as H. halys. In forest margins surrounding peach, pecan, and interplanted peach-pecan orchards, we monitored H. halys populations with pheromone-baited traps, mist-netted birds, and collected avian fecal samples for molecular gut content analysis. We screened 257 fecal samples from 19 bird species for the presence of H. halys DNA to determine whether birds provide the biological control of this pest. Overall, we found evidence that four birds from three species consumed H. halys, including Northern cardinal (Cardinalis cardinalisis), Tufted titmouse (Baeolophus bicolor), and Carolina wren (Thryothorus ludovicianus). Halyomorpha halys captured in traps increased over time but did not vary by orchard type. Although incidence of predation was low, this may be an underestimate as a result of our current avian fecal sampling methodology. Because birds are members of the broader food web, future studies are needed to understand avian ecosystem services, especially in terms of pest control, including H. halys and other pest species.

Geographically consistent hybridization dynamics between the Black-crested and Tufted titmouse with evidence of hybrid zone expansion

Abstract We studied hybridization between the Black-crested and Tufted titmouse across two geographically distinct transects that differ in the timing of secondary contact by hundreds to thousands of years. We found that hybridization patterns correspond to localized hybrid swarms and that the titmouse hybrid zone is likely slowly expanding over time, a product of short post-natal dispersal distances coupled with weak or absent selection against admixture. We show the southern part of the hybrid zone located in Texas is four times wider than the northern region of hybridization in Oklahoma, which is likely due to geographic differences in hybrid zone age. Despite differences in width, most individuals in both transects are advanced-generation hybrids and backcrosses, suggesting geographically consistent hybridization dynamics. We documented a strong correlation between genotypes and plumage index, suggesting that hybridization has not yet resulted in the decoupling of plumage and genome-wide ancestry as observed in some other avian hybrid zones. Although our results suggest the ongoing expansion of the hybrid zone, the rate of expansion appears to be slow, on the scale of tens of meters a year, and it will likely take hundreds of thousands to millions of years before homogenization of the parental populations. While we did not find support for partial reproductive isolation in the hybrid zone itself, there is the possibility that ecological or sexual selection limits introgression into allopatric regions. Broadly, the results of our study highlight the value of multiple, geographically distant, transects across a hybrid zone for assessing the evolutionary dynamics of hybridizing lineages.

Concentrations of Manganese in Tufted Titmouse Feathers near Metal Processing Plants

Human industry and land use has led to the anthropogenic release of manganese (Mn) into the air and soil near manufacturing centers. Overexposure to Mn can cause considerable health problems in birds. We studied whether the concentration of Mn in bird feathers correlates with the distance to point sources of Mn air emissions. Feathers were collected from Tufted Titmice (Baeolophus bicolor) at two sites in western Pennsylvania, USA. One site was in proximity (0.3 km) to a steel plant with documented Mn releases, and the other site was in a different town about 4.0 km away from other steel plants with documented Mn releases. Using the microwave plasma–atomic emission spectrometer (MP–AES), we found that tail feathers collected from nearest to a steel plant had a significantly higher concentration of Mn compared to the samples from the site further from the emission source. A body mass index was calculated for each set of birds; however, the indices did not vary significantly. This is the first published study of Mn sequestration in Tufted Titmouse feathers. This study develops our general understanding of the potential use of bird feathers as non-invasive bioindicators of environmental metal exposure.

Foraging Intensity of Tufted Titmice (Baeolophus bicolor) in Response to Feed Type and Extreme Winter Conditions

Foraging-intensity models predict that consumers balance the costs of environmental risk factors, food availability, predation, and starvation to determine the optimal foraging rate response. To test the assumptions of these ecological models in situ, we used radio frequency identification (RFID) to monitor the foraging behavior of 5 individual Baeolophus bicolor (Tufted Titmice) at bird feeders from January to February 2021. We used a Poisson generalized linear mixed effects model to determine how temperature and seed type impacted the number of daily feeder visits, while controlling for feeder and individual ID. We documented increased foraging frequency for Titmice in colder, snowier conditions. Titmice also exhibited a strong preference for whole black oil sunflower seeds (with an intact outer shell), in contrast to sunflower hearts (with the shell removed).

Social dynamics of core members in mixed-species bird flocks change across a gradient of foraging habitat quality.

Social associations within mixed-species bird flocks can promote information flow about food availability and provide predator avoidance benefits. The relationship between flocking propensity, foraging habitat quality, and interspecific competition can be altered by human-induced habitat degradation. Here we take a close look at sociality within two ecologically important flock-leader (core) species, the Carolina chickadee (Poecile carolinensis) and tufted titmouse (Baeolophus bicolor), to better understand how degradation of foraging habitat quality affects mixed-species flocking dynamics. We compared interactions of free ranging wild birds across a gradient of foraging habitat quality in three managed forest remnants. Specifically, we examined aspects of the social network at each site, including network density, modularity, and species assortativity. Differences in the social networks between each end of our habitat gradient suggest that elevated levels of interspecific association are more valuable in the habitat with low quality foraging conditions. This conclusion is supported by two additional findings: First, foraging height for the subordinate Carolina chickadee relative to the tufted titmouse decreased with an increase in the number of satellite species in the most disturbed site but not in the other two sites. Second, the chickadee gargle call rate, an acoustic signal emitted during agonistic encounters between conspecifics, was relatively higher at the high-quality site. Collectively, these results suggest an increase in heterospecific associations increases the value of cross-species information flow in degraded habitats.

What the pluck? The theft of mammal hair by birds is an overlooked but common behavior with fitness implications.

Avian species across diverse lineages collect and incorporate mammalian hair into their nests (Tóth 2008). This widespread behavior can be adaptive, as hair, fur or wool insulates nests and so enhances nestling survival and recruitment in colder climates (Hilton et al. 2004, Mainwaring et al. 2014, Järvinen and Brommer 2020, Deeming et al. 2020; reviewed in Perez et al. 2020). How birds obtain mammal hair for their nests, however, is an open question. A common assumption is that birds gather mammal hair that has been shed into the environment or from carcasses (e.g. Tóth 2008), although anecdotal accounts exist in the scientific literature of birds plucking hair directly from live mammals. Here, we (H.S.P., Z.S.S. and J.D.B.) document and report our observation of a Tufted Titmouse (Baeolophus bicolor) exhibiting a behavior that we are terming "kleptotrichy" (from Greek "klepto-" - to steal + "trich-" - hair), after "kleptoptily" or feather theft described by Whitney (2007).

Homeward bound: canopy cover and species identity influence non-breeding season homing success and speed in forest birds

BackgroundEfficient and safe movement is fundamental for wild birds to thrive in their environments. For arboreal forest animals, especially birds, canopy cover has a large impact on birds’ daily movements and is a crucial component of conservation strategies seeking to retain avian population in disturbed or urban habitats.MethodsWe translocated woodland bird species utilizing different forest strata during two non-breeding seasons in Gainesville, FL, USA. We used linear model and generalized linear model to examine the effects of canopy cover and species identity on homing success and speed.ResultsAmong our study species of Tufted Titmouse (Baeolophus bicolor), Carolina Chickadee (Poecile carolinensis), and Northern Cardinal (Cardinalis cardinalis), we found that Carolina Chickadees and Tufted Titmice were more likely to return than Northern Cardinals. Among birds that successfully returned, homing speed is significantly affected by forest canopy cover and species identity (titmice had higher homing speed than cardinals). Birds return much faster in landscape with higher canopy cover.ConclusionsThis study presented evidence of species identity’s effect on homing success and speed in common feeder bird species in Southeast US and provided further evidence that bird movements in the suburban land cover are constrained by low canopy cover.

A community context for aggression? Multi-species audience effects on territorial aggression in two species of Paridae.

Territorial aggression in birds is widely observed and is commonly linked to sex, age, body size, physiology, seasonal cues, food resource, urbanization, and a variety of social contexts including conspecific audience effects. However, little is known about the heterospecific audience effects on territorial aggression.Here, we address an emerging idea that heterospecific audience effects may be pervasive influences in the social lives of free‐living birds. We tested the hypothesis that the composition, number, and relative body size of heterospecific audiences observing an aggressive contest will influence the response probability and intensity of aggression displayed.We subjected two Paridae species, tufted titmouse (TUTI, Baeolophus bicolor) and Carolina chickadee (CACH, Poecile carolinensis), to playbacks of aggressive calls during a breeding season in north‐central Florida. At widely spaced playback sites (N = 134) in woodland habitats, we characterized the makeup of heterospecific audiences, aggression type (intra vs. interspecific territoriality), local population density, and various environmental factors (tree density, wind speed, and noise level) that are likely to influence territorial aggression.We found that the presence of heterospecific audiences increased TUTI aggression levels and that both parids were more likely to respond to playback stimuli when their audiences had higher heterospecific diversity (more heterospecific individuals and species). We also found TUTI were more likely to respond when CACH were present but not vice versa.In conclusion, we found evidence that heterospecific audiences significantly influenced the metrics of territorial aggression of free‐living animals and we suggest that the definition of audience effects on the behavior of free‐living animals be expanded to incorporate heterospecific audiences.

Variable or atypical? Comparing unusual songs of the Tufted Titmouse with a citizen-science database

The Tufted Titmouse Baeolophus bicolor is well known for its stereotyped “peter-peter-peter” song, but anecdotal reports of its song’s diversity also exist. We identified an atypical song in East-Central Illinois, USA, that has persisted for several years in the same vicinity. We used a spectrogram-based comparison metric to assess song similarity scores against con- and heterospecific songs from a citizen-science database. The results suggest that these novel songs may truly be outliers beyond the Tufted Titmouse’s local range of species-typical song variability. Our study also illustrates a methodological approach regarding the assessment of other, potentially atypical vocalizations recorded from birds.

Experimental manipulation of mixed-species flocks reveals heterospecific audience effects on calling

Animal signallers are subject to audience effects when they alter communication due to changes in the presence or characteristics of receivers. Studies aimed at understanding audience effects have typically examined effects of conspecific audiences on signaller communication. Less work has focused on heterospecific audiences, which present an important avenue of research for species that participate in mixed-species groups. Here we experimentally tested mixed-species flocks of Carolina chickadees, Poecile carolinensis, and tufted titmice, Baeolophus bicolor, for conspecific and heterospecific audience effects. Birds were trapped from naturally occurring flocks and held in seminatural outdoor aviaries, where we recorded calling. We found that chickadees and titmice were sensitive to the number of conspecifics in flocks when communicating via ‘chick-a-dee’ calls, which are social cohesion calls produced by both species. Chickadees also were sensitive to the number of titmice in flocks, but chick-a-dee calling behaviour in titmice did not differ with regard to the number of chickadees in flocks. Furthermore, when subject to playbacks of simulated risk, chickadees and titmice produced more chick-a-dee calls when more titmice were in a flock. After these playbacks, chickadees produced fewer chick-a-dee calls with increasing numbers of conspecifics in flocks, whereas titmice produced more chick-a-dee calls with increasing numbers of conspecifics in flocks. These results suggest that chickadees and titmice are sensitive to social factors within their mixed-species flocks when communicating, and that chickadees appear more sensitive to heterospecific presence than do titmice. We suggest this is due to the dominance status of the species in these flocks, where titmice are typically dominant over chickadees.

Tufted Titmice (Baeolophus bicolor) rear a mixed brood to apparent fledging in northeastern Arkansas

Interspecific nest usurpation, a relatively common nesting strategy, is the act of one bird species taking over the nesting site of another species. Although used by secondary cavity-nesters, nest usurpation has never been reported for the Tufted Titmouse (Baeolophus bicolor). In this paper, we document a mixed brood successfully reared by Tufted Titmice after usurpation of an Eastern Bluebird (Sialia sialis) nest in northeastern Arkansas during the 2019 breeding season. We monitored the focal nest box every 3–4 d from the first sign of nest building on 23 March 2019 until chick fledging by 12 May 2019. The Tufted Titmice took over the nest at the time the Eastern Bluebird had laid its fourth egg in the nest. By the end of the nesting period, the Tufted Titmice fledged 2 of their own chicks and 3 Eastern Bluebird chicks. Such successful rearing of non-conspecific nestlings is rare and we discuss factors that may have allowed it.

Traffic noise and responses to a simulated approaching avian predator in mixed‐species flocks of chickadees, titmice, and nuthatches

AbstractTraffic noise likely reaches a wide range of species and populations throughout the world, but we still know relatively little about how it affects anti‐predator behavior of populations. We tested for possible effects of traffic noise on responses to predator acoustic cues in Carolina chickadees (Poecile carolinensis), tufted titmice (Baeolophus bicolor), and white‐breasted nuthatches (Sitta carolinensis) near 14 independent feeding stations in eastern Tennessee. We compared anti‐predator calling and seed‐taking behavior in response to playbacks of predator stimuli (screech owl calls) at sites naturally exposed to traffic noise and at sites that faced relatively little traffic noise. The screech owl call playback was designed to simulate the approach of this dangerous predator to a feeder being used by these small songbirds. We found that chickadees responded consistently to the owl stimuli across different levels of traffic noise. However, titmice, and nuthatches exhibited different behavioral responses to the predator stimulus, suggesting that traffic noise masked these low‐frequency predator calls. Overall, chickadees and nuthatches showed the broadest anti‐predator behavioral responses in comparison to titmice, corroborating earlier published work with an Indiana population. Finally, populations exposed to traffic noise overall seemed less able to detect predator cues potentially masked by that noise, and future work will need to assess likely seasonal variation in these responses as well as species‐level variation in anti‐predator responses in mixed‐species groups.

Foraging ecology drives social information reliance in an avian eavesdropping community.

Vertebrates obtain social information about predation risk by eavesdropping on the alarm calls of sympatric species. In the Holarctic, birds in the family Paridae function as sentinel species; however, factors shaping eavesdroppers' reliance on their alarm calls are unknown. We compared three hypothesized drivers of eavesdropper reliance: (a) foraging ecology, (b) degree of sociality, and (c) call relevance (caller‐to‐eavesdropper body‐size difference). In a rigorous causal‐comparative design, we presented Tufted Titmouse (Baeolophus bicolor) alarm calls to 242 individuals of 31 ecologically diverse bird species in Florida forests and recorded presence/absence and type (diving for cover or freezing in place) of response. Playback response was near universal, as individuals responded to 87% of presentations (N = 211). As an exception to this trend, the sit‐and‐wait flycatcher Eastern Phoebe (Sayornis phoebe) represented 48% of the nonresponses. We tested 12 predictor variables representing measures relevant to the three hypothesized drivers, distance to playback speaker, and vulnerability at time of playback (eavesdropper's microhabitat when alarm call is detected). Using model‐averaged generalized linear models, we determined that foraging ecology best predicted playback response, with aerial foragers responding less often. Foraging ecology (distance from trunk) and microhabitat occupied during playback (distance to escape cover) best predicted escape behavior type. We encountered a sparsity of sit‐and‐wait flycatchers (3 spp.), yet their contrasting responses relative to other foraging behaviors clearly identified foraging ecology as a driver of species‐specific antipredator escape behavior. Our findings align well with known links between the exceptional visual acuity and other phenotypic traits of flycatchers that allow them to rely more heavily on personal rather than social information while foraging. Our results suggest that foraging ecology drives species‐specific antipredator behavior based on the availability and type of escape cover.

Estimation of growth regulation in natural populations by extended family of growth curve models with fractional order derivative: Case studies from the global population dynamics database

Estimating the trend in population time series data using growth curve models is a central idea in population ecology. Several models, mainly governed by differential or difference equations, have been applied to real data sets to identify general growth pattern and make predictions. In this article, we analyze ecological time series data by fitting mathematical models governed by fractional differential equations (FDE). The order of the FDE (α) is used to quantify the evidence of memory in the population processes. The application of FDE is exemplified by analyzing time series data on two bird species Phalacrocorax carbo (Great cormorant) and Parus bicolor (Tufted titmouse) and two mammal species Castor canadensis (Beaver) and Ursus americanus (American black bear) extracted from the global population dynamics database. Five different population growth models were fitted to these data; density-independent exponential, negative density-dependent logistic and θ-logistic model, positive density-dependent exponential Allee and strong Allee model. Both ordinary and fractional derivative representations of these models were fitted to the time series data. Markov chain Monte Carlo (MCMC) method was used to estimate the model parameters and Akaike information criterion was used to select the best model. By estimating the return rate for each of the time series, we have shown that populations governed by FDE with a small value of α (high level of memory) return to the stable equilibrium faster. This demonstrates a synergistic interplay between memory and stability in natural populations.

Flocking propensity by satellites, but not core members of mixed-species flocks, increases when individuals experience energetic deficits in a poor-quality foraging habitat.

Mixed-species bird flocks are complex social systems comprising core and satellite members. Flocking species are sensitive to habitat disturbance, but we are only beginning to understand how species-specific responses to habitat disturbance affect interspecific associations in these flocks. Here we demonstrate the effects of human-induced habitat disturbance on flocking species’ behavior, demography, and individual condition within a remnant network of temperate deciduous forest patches in Indiana, USA. Specifically, we characterized the following properties of two core species, Carolina chickadees (Poecile carolinensis) and tufted titmice (Baeolophus bicolor), across a secondary-forest disturbance gradient: foraging time budgets, home range size, fat scores, fledgling counts, survival rates, and abundance. We also report fat scores for two satellite species that flock with the core study species: white-breasted nuthatches (Sitta carolinensis) and downy woodpeckers (Dryobates pubescens). Finally, we assess mixed-species flock sizes and composition, in addition to avian predator call rates, across the disturbance gradient. Foraging time budgets and home range size were highest and fat scores were lowest for core species in the most-disturbed site. Fat scores of two satellite species followed the same pattern. Additionally, the number of tufted titmice fledglings and winter survival rate of Carolina chickadees were lowest at the most-disturbed site. These results suggest that core species in the most-disturbed site experienced energetic deficits. Moreover, cumulative calling rate of raptors was lowest at the most-disturbed site, and none of the individual raptor species call rates were higher at the most-disturbed site—suggesting that perception of predation risk does not contribute to these patterns. Surprisingly, the satellites continued associating with mixed species flocks through the breeding season at the most-disturbed site. Total flock size and interspecific association patterns were otherwise consistent across the gradient. The fact that satellites continued to flock with core species during the breeding season suggests foraging niche expansion resulting from mixed-species flocking is important in disturbed sites even beyond the winter season. Our study reveals mechanisms underlying flock composition of birds surviving in remnant forest and links the mechanisms to degradation of foraging habitat. These findings offer important insight into the relative impact potential of forest disturbance on mixed-species flocks in the North Temperate Zone.