Fig. 7

Expression of the messaging ‘syndrome’ follows a stereotypical developmental trajectory. a-b Active attraction of workers to the queen and contact rates between the queen and workers as a function of worker age. Points represent colony means, and point colours indicate colony identity. Solid lines indicate general additive models, fitted to the colony means. c-d Community affiliations in the daily contact networks as a function of age. e-f Spreading roles in the daily relay network according to age. Out-degree \(k_{out}\) represents the number of nestmates an individual donates queen pheromone to, and degree difference \(\Delta k\) represents the out-degree minus the in-degree (number of nestmates an individual receives queen pheromone from). g-h Dimension reduction reveals that workers follow stereotypical developmental trajectories as they age. g Principal component analysis of the six behaviours shown in panels a-f. Background points represent a given individual on a given day. Foreground points show the mean for a given age cohort on a given day. Point colours represent colony identities. h Blowup of the rectangular area in g. Coloured arrows show developmental trajectories for each colony. The black arrow shows the ‘global’ trajectory. Coloured points show the inflection point for each colony. Points marked with an ‘x’ indicate the 3–5 day-old workers, which contribute most to feeding the queen and raising new queens. i-k The daily contact network (i-j) and the daily pheromone relay network (k) for colony 18 on 29/8/2016. All networks used the same layout, obtained by applying a force-directed layout algorithm to the contact network. Nodes represent individual bees, and the queen is indicated by the white node. Node size indicates age. In (i-j), edge thickness represents the number of pairwise encounters. In (k) edge weight & direction indicates pheromone flow. In (i), the nodes are coloured according to the community scores. The network is partitioned into three overlapping communities; a foraging community consisting mainly of older workers (red nodes), and two nurse communities consisting mainly of younger workers (blue and green nodes). In (j) nodes are coloured according to the nurse bridging score, \(H^\prime (N_A,N_B)\). Bees positioned at the overlap of the two nurse communities have high bridging scores. In (k) nodes are coloured according to the out degree. l The coarse-grained flow network. Node labels indicate the age in days. Weighted and directed edges indicate the net pheromone flow between cohorts. The donation hierarchy is indicated by the vertical positions of each node, as defined by the dominance ranks