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.

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.

Individual variation in foraging behavior reveals a trade-off between flexibility and performance of a top predator.

There is increasing evidence that behavioral flexibility is associated with the ability to adaptively respond to environmental change. Flexibility can be advantageous in some contexts such as exploiting novel resources, but it may come at a cost of accuracy or performance in ecologically relevant tasks, such as foraging. Such trade-offs may, in part, explain why individuals within a species are not equally flexible. Here, we conducted a reversal learning task and predation experiment on a top fish predator, the Northern pike (Esox lucius), to examine individual variation in flexibility and test the hypothesis that an individual's behavioral flexibility is negatively related with its foraging performance. Pikes were trained to receive a food reward from either a red or blue cup and then the color of the rewarded cup was reversed. We found that pike improved over time in how quickly they oriented to the rewarded cup, but there was a bias toward the color red. Moreover, there was substantial variation among individuals in their ability to overcome this red bias and switch from an unrewarded red cup to the rewarded blue cup, which we interpret as consistent variation among individuals in behavioral flexibility. Furthermore, individual differences in behavioral flexibility were negatively associated with foraging performance on ecologically relevant stickleback prey. Our data indicate that individuals cannot be both behaviorally flexible and efficient predators, suggesting a trade-off between these two traits.

Maternal predator-exposure has lifelong consequences for offspring learning in threespined sticklebacks.

Learning is an important form of phenotypic plasticity that allows organisms to adjust their behaviour to the environment. An individual's learning performance can be affected by its mother's environment. For example, mothers exposed to stressors, such as restraint and forced swimming, often produce offspring with impaired learning performance. However, it is unclear whether there are maternal effects on offspring learning when mothers are exposed to ecologically relevant stressors, such as predation risk. Here, we examined whether maternal predator-exposure affects adult offsprings' learning of a discrimination task in threespined sticklebacks (Gasterosteus aculeatus). Mothers were either repeatedly chased by a model predator (predator-exposed) or not (unexposed) while producing eggs. Performance of adult offspring from predator-exposed and unexposed mothers was assessed in a discrimination task that paired a particular coloured chamber with a food reward. Following training, all offspring learned the colour-association, but offspring of predator-exposed mothers located the food reward more slowly than offspring of unexposed mothers. This pattern was not driven by initial differences in exploratory behaviour. These results demonstrate that an ecologically relevant stressor (predation risk) can induce maternal effects on offspring learning, and perhaps behavioural plasticity more generally, that last into adulthood.