Friends of friends: are indirect connections in social networks important to animal behaviour?

Friend of a friend relationships, or the indirect connections between people, influence our health, well-being, financial success and reproductive output. As with humans, social behaviours in other animals often occur within a broad interconnected network of social ties. Yet studies of animal social behaviour tend to focus on associations between pairs of individuals. With the increase in popularity of social network analysis, researchers have started to look beyond the dyad to examine the role of indirect connections in animal societies. Here, I provide an overview of the new knowledge that has been uncovered by these studies. I focus on research that has addressed both the causes of social behaviours, i.e. the cognitive and genetic basis of indirect connections, as well as their consequences, i.e. the impact of indirect connections on social cohesion, information transfer, cultural practices and fitness. From these studies, it is apparent that indirect connections play an important role in animal behaviour, although future research is needed to clarify their contribution.