Honeybees are renowned for their perfectly hexagonal honeycomb, hailed as the pinnacle of biological architecture for its ability to maximize storage area while minimizing building material. However, in natural nests, workers must regularly transition between different cell sizes, merge inconsistent combs, and optimize construction in constrained geometries. These spatial obstacles pose challenges to workers building perfect hexagons, but it is unknown to what extent workers act as architects versus simple automatons during these irregular building scenarios. Using automated image analysis to extract the irregularities in natural comb building, we show that some building configurations are more difficult for the bees than others, and that workers overcome these challenges using a combination of building techniques, such as: intermediate-sized cells, regular motifs of irregular shapes, and gradual modifications of cell tilt. Remarkably, by anticipating these building challenges, workers achieve high-quality merges using limited local sensing, on par with analytical models that require global optimization. Unlike automatons building perfectly replicated hexagons, these building irregularities showcase the active role that workers take in shaping their nest and the true architectural abilities of honeybees.
Keywords: collective behavior; nest construction; optimization; spatial structure; superorganism.