Egalitarian mixed-species bird groups enhance winter survival of subordinate group members but only in high-quality forests

Abstract

Only dominant individuals have unrestricted access to contested resources in group-living animals. In birds, subordinates with restricted access to resources may respond to intragroup contests by acquiring extra body reserves to avoid periods of food shortage. In turn, higher body mass reduces agility and increases predation and mortality risk to subordinates. Birds often live in hierarchically organized mixed-species groups, in which heterospecific individuals are considered to substitute for conspecifics as protection against predators at a significantly reduced competition cost. Crested tits (Lophophanes cristatus) and willow tits (Poecile montanus) form mixed-species groups during the non-reproductive season that typically exhibit a nearly linear dominance hierarchy (‘despotic’ social structure) in which the highest ranking male willow tit is fourth in the overall hierarchy after the dominant male, female and subordinate juvenile crested tit, respectively. Much less frequently, ‘egalitarian’ dominance structures occur in which the adult willow tits rank second and the hierarchy is less steep, or linear. We present a rare long-term data set in which egalitarian flocks are common enough to assess the consequences of this simple change in hierarchy structure as well as a potential driver of the pattern. A comparison of individuals in the despotic mixed-species groups revealed a strong negative correlation between subcutaneous fat stores and dominance rank in the interspecific dominance hierarchy, whereas in egalitarian groups, subordinate willow tits had significantly lower fat reserves and they foraged in safer parts of the canopy than willow tits in despotic groups. Moreover, egalitarian groups exhibited markedly less within-group aggression, higher group cohesion and improved winter survival in both tit species. However, winter survival of birds in egalitarian groups was impaired relative to despotic groups in forests recently affected by industrial forestry. This suggests that the more egalitarian bird societies may best be adapted to less-disturbed environments.