Fig. 1: Spatial organisation in four species of social insects.

a–d One-hour trajectories of all workers in an example colony. The nest interior is divided into discrete sites defined by a hexagonal grid. The trajectory of a single focal individual is shown in red. e–h Corresponding site-visit networks. Edges connect individuals (upper layer) to the sites they visit (bottom layer). Edge widths are proportional to the number of visits. The module scores for each node are indicated by the coloured bars (Nurse (N): cyan; Intermediate (I): black; Peripheral (P): magenta; Forager (F): yellow). In the honeybee colony, two different shades of cyan are used to depict the two distinct nurse modules. The connections of the focal individual are shown in red. i–l Map of the nest floor. Sites are coloured according to the linear combination of CMYK colours for each module, weighted by its module scores; thus a site with a forager score of 1 (‘pure’ forager site) is shown in yellow (top right corner in the colour key), while a site with a forager score of 0.5 and a peripheral score of 0.5 is shown in orange (middle of top row in the colour key). In the honeybee colony, two different shades of cyan are used to depict the two distinct nurse modules. Solid black lines indicate the borders of the broodnest in A. mellifera, and the outline the brood pile in the ants. The circle indicates the nest entrance. White grid cells correspond to unvisited sites (typically occupied by clusters of brood in the ants, or wax pillars in the bees). m–p Heatmaps showing the module score diversity of all visitors to each site (‘Visitor diversity’). The brood and the area near the nest entrance are associated with low visitor diversity (i.e., low mixing between individuals from different modules), whilst the intermediate area is associated with high visitor diversity (i.e., high mixing between individuals from different modules). White grid cells correspond to unvisited sites. Source data are provided as a source data file.