Social, spatial, and temporal organization in a complex insect society.

High-density living is often associated with high disease risk due to density-dependent epidemic spread. Despite being paragons of high-density living, the social insects have largely decoupled the association with density-dependent epidemics. It is hypothesized that this is accomplished through prophylactic and inducible defenses termed 'collective immunity'. Here we characterise segregation of carpenter ants that would be most likely to encounter infectious agents (i.e. foragers) using integrated social, spatial, and temporal analyses. Importantly, we do this in the absence of disease to establish baseline colony organization. Behavioural and social network analyses show that active foragers engage in more trophallaxis interactions than their nest worker and queen counterparts and occupy greater area within the nest. When the temporal ordering of social interactions is taken into account, active foragers and inactive foragers are not observed to interact with the queen in ways that could lead to the meaningful transfer of disease. Furthermore, theoretical resource spread analyses show that such temporal segregation does not appear to impact the colony-wide flow of food. This study provides an understanding of a complex society's organization in the absence of disease that will serve as a null model for future studies in which disease is explicitly introduced.