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Honey bees (Apis mellifera L.) localize the queen and aggregate into a swarm by forming a collective scenting network to directionally propagate volatile pheromone signals. Previous experiments show the robustness of this communication strategy in the presence of physical obstacles that partially block pheromone flow and the path to the queen. Specifically, there is a delay in the formation of the scenting network and aggregation compared to a simple environment without perturbations. To better understand the effect of obstacles beyond temporal dynamics, we use the experimental results as inspiration to explore how the behavioral parameter space of collective scenting responds to obstacle. We extend an agent-based model previously developed for a simple environment to account for the presence of physical obstacles. We study how individual agents with simple behavioral rules for scenting and following concentration gradients can give rise to collective localization and swarming. We show that the bees are capable of navigating the more complex environment with a physical obstacle to localize the queen and aggregate around her, but their range of behavioral parameters is more limited and less flexible as a result of the spatial density heterogeneity in the bees imposed by the obstacle. 

Honeybee swarms are a landmark example of collective behavior. To become a coherent swarm, bees locate their queen by tracking her pheromones. But how can distant individuals exploit these chemical signals, which decay rapidly in space and time? Here, we combine a behavioral assay with the machine vision detection of organism location and scenting (pheromone propagation via wing fanning) behavior to track the search and aggregation dynamics of the honeybee Apis mellifera L. We find that bees collectively create a scenting-mediated communication network by arranging in a specific spatial distribution where there is a characteristic distance between individuals and directional signaling away from the queen. To better understand such a flow-mediated directional communication strategy, we developed an agent-based model where bee agents obeying simple, local behavioral rules exist in a flow environment in which the chemical signals diffuse and decay. Our model serves as a guide to exploring how physical parameters affect the collective scenting behavior and shows that increased directional bias in scenting leads to a more efficient aggregation process that avoids local equilibrium configurations of isotropic (nondirectional and axisymmetric) communication, such as small bee clusters that persist throughout the simulation. Our results highlight an example of extended classical stigmergy: Rather than depositing static information in the environment, individual bees locally sense and globally manipulate the physical fields of chemical concentration and airflow.