What constitutes "social complexity" and "social intelligence" in birds? Lessons from ravens

In the last decades, the assumption that complex social life is cognitively challenging, and thus can drive mental evolution, has received much support from empirical studies in nonhuman primates. While extending the scope to other mammals and birds, different views have been adopted on what constitutes social complexity and which specific cognitive skills are selected for. Notably, many avian species form open groups as non-breeders (i.e., seasonally and before sexual maturity) that have been largely ignored as potential sources of social complexity. Reviewing 30years of research on ravens, we illustrate the socio-ecological conditions faced by these birds as non-breeders and discuss how these relate to their socio-cognitive skills. We argue that the non-breeding period is key to understand raven social life and, to a larger extent, avian social life in general. We furthermore emphasize how the combination of the large-scale perspective (defining social system components: e.g., social organization, mating system) and the individual-scale perspective on social systems allows to better capture the complete set of social challenges experienced by individuals throughout their life, ultimately resulting on a more comprehensive understanding of species' social complexity.