Flexible foraging shapes the topology of plant-pollinator interaction networks

In plant-pollinator networks, foraging choices by pollinators help form the connecting links between species. Flexible foraging should therefore play an important role in defining network topology. Factors such as morphological trait complementarity limit a pollinator's pool of potential floral resources, but which potential resource species are actually utilized at a location depends on local environmental and ecological context. Pollinators can be highly flexible foragers, but the effect of this flexibility on network topology remains unclear. To understand how flexible foraging affects network topology, we examined differences between sets of locally realized interactions and corresponding sets of potential interactions within 25 weighted plant-pollinator networks in two different regions of the United States. We examined two possible mechanisms for flexible foraging effects on realized networks: (1) preferential targeting of higher-density plant resources, which should increase network nestedness, and (2) context-dependent resource partitioning driven by interspecific competition, which should increase modularity and complementary specialization. We found that flexible foraging has strong effects on realized network topology. Realized connectance was much lower than connectance based on potential interactions, indicating a local narrowing of diet breadth. Moreover, the foraging choices pollinators made, which particular plant species to visit and at what rates, resulted in networks that were significantly less nested and significantly more modular and specialized than their corresponding networks of potential interactions. Preferentially foraging on locally abundant resources was not a strong driver of the realization of potential interactions. However, the degree of modularity and complementary specialization both increased with the number of competing pollinator species and with niche availability. We therefore conclude that flexible foraging affects realized network topology more strongly through resource partitioning than through focusing on high-density resources.