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, ) and Carolina chickadee (CACH, ), to playbacks of aggressive calls during a breeding season in north-central Florida. At widely spaced playback sites ( = 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.