Indirectly connected: simple social differences can explain the causes and apparent consequences of complex social network positions.

Animal societies are often structurally complex. How individuals are positioned within the wider social network (i.e. their indirect social connections) has been shown to be repeatable, heritable and related to key life-history variables. Yet, there remains a general lack of understanding surrounding how complex network positions arise, whether they indicate active multifaceted social decisions by individuals, and how natural selection could act on this variation. We use simulations to assess how variation in simple social association rules between individuals can determine their positions within emerging social networks. Our results show that metrics of individuals' indirect connections can be more strongly related to underlying simple social differences than metrics of their dyadic connections. External influences causing network noise (typical of animal social networks) generally inflated these differences. The findings demonstrate that relationships between complex network positions and other behaviours or fitness components do not provide sufficient evidence for the presence, or importance, of complex social behaviours, even if direct network metrics provide less explanatory power than indirect ones. Interestingly however, a plausible and straightforward heritable basis for complex network positions can arise from simple social differences, which in turn creates potential for selection to act on indirect connections.