A model of resource partitioning between foraging bees based on positive and negative associations

Central place foraging pollinators tend to develop multi-destination routes (traplines) to visit patchily distributed plant resources. While the formation of traplines by individual pollinators has been studied in details, how populations of foragers exploit resources in a common area is an open question, difficult to address experimentally. We explored conditions for the emergence of resource partitioning among traplining bees using agent-based models built from experimental data of bumblebees foraging on artificial flowers. In the models, bees learn to develop routes as a consequence of feedback loops that change their probabilities of moving between flowers. While a positive reinforcement of movements leading to rewarding flowers is sufficient for the emergence of resource partitioning when flowers are evenly distributed, addition of negative reinforcement of movements leading to unrewarding flowers is necessary when flowers are patchily distributed. In environments with more complex spatial structure, the negative experiences of individual bees on flowers favour spatial segregation and efficient collective foraging.