Modelling foraging strategies of honey bees as agents in a dynamic landscape representation

Abstract

Introduction: Both intrinsic colony mechanisms and external environmental variables affect the honey bee colony development rates and response and a key aspect of this is the use of resources within the landscape by honey bees. Although several models have been developed to explore the foraging behaviour of bees, none fully considered the spatial and temporal dynamics of landscape resources and the role of the colony in the process. Methodology: Here, we introduce a new honey bee foraging model being developed as a part of the ApisRAM honey bee colony model. Based on agent-based modelling, we used a dynamic ALMaSS landscape model enhanced with floral resource modelling to assess the impacts of weather conditions and resource availability on the possible foraging behaviour of honey bees. Several possible mechanisms (defined, based on honey bee traits) for scouting and foraging were investigated, separately for nectar and pollen collection, including prioritising foraging polygons for nectar foraging according to their distance to the colony, the quality or the energetic efficiency and, for pollen foraging, according to their distance to the colony and pollen quantity. Results: If model foraging bees prioritised the polygons, based on their distance from the colony, the number of unsuccessful flights increased compared to other tested strategies and the total amount of sugar collected showed a high variability. Contrary to expectations, the energetic efficiency strategy did not provide the colony with the highest amount of sugar. Overall, the tested strategies provide different outcomes on the collection of resources, the number of performed flights and their success rate, evidencing that the model's outcome at the colony level arises from the individual types of behaviour. Conclusions and Relevance: Variability in the mass of collected nectar and pollen was found mostly when scout bees applied the distance strategy. This higher variability in sugar collection means that model bees were not able to find the most profitable foraging sites at the landscape level, but only at the local level. Other strategies showed less dependence on the surrounding landscape (i.e. quality or random), but it comes at a cost (i.e. lower production for both nectar and pollen collection). These outputs help us evaluate which strategies could be used for future model development and confirm the models' ability to create dynamic responses. These responses at the colony level were only possible thanks to the implementation of a dynamic landscape model and dynamic spatiotemporal resource model, as well as implementing a social communication mechanism for bees to share information about the resources. Plant nectar production and quality information is essential to predict honey bee foraging distribution. In future model development, the implementation of pollen quality should also be explored to evaluate if it influences the overall pollen collection.