Abstract
Flowers cluster at various spatial scales, so pollinators use information from multiple scales when foraging in natural plant populations. Little is known about the effects of interactions between scales or their relative strength. We examined bumblebee foraging behaviour in a natural population of Salvia nipponica in 10 and 7 patches in 2019 and 2020, respectively. We recorded within-patch factors (display size of racemes and local open raceme densities) and patch-level factors (patch size and distance from the nearest patch) and analysed their relationships with pollinator behaviour. The numbers of visits per raceme and flower were mainly affected by the interaction of patch size and raceme density; they were higher in locations with lower raceme density in larger patches. The ratio of flowers visited to all open flowers in a raceme during a raceme visit, which relates to a bumblebee's choice to leave a raceme, was mainly affected by the interaction of display size and local open raceme density; in 2019 it was higher in racemes with smaller display sizes, while in 2020 the strength and direction of the relationship depended on the open raceme density. These results suggest that pollinators relied on the sizes of flower clusters at different spatial scales when visiting and leaving racemes and adjusted their responses to the sizes of flower clusters depending on the distances between clusters. Therefore, it is important to evaluate factors at various spatial scales and their interactions to fully understand pollinator behaviour in natural plant populations.