ABSTRACT
Foraging behavior of pollinators is shaped by, among other factors, the conflict between maximizing resource intake and minimizing predation risk; yet, empirical studies quantifying variation in both forces are rare, compared to those investigating each separately. Here, we discuss the importance of simultaneously assessing bottom-up and top-down forces in the study of plant-pollinator interactions, and propose a conceptual and testable graphical hypothesis for pollinator foraging behavior and plant fitness outcomes as a function of varying floral rewards and predation risk. In low predation risk scenarios, no noticeable changes in pollinator foraging behavior are expected, with reward levels affecting only the activity threshold. However, as predation risk increases we propose that there is a decrease in foraging behavior, with a steeper decline as plants are more rewarding and profitable. Lastly, in high predation risk scenarios, we expect foraging to approach zero, regardless of floral rewards. Thus, we propose that pollinator foraging behavior follows an inverse S-shape curve, with more pronounced changes in foraging activity at intermediate levels of predation risk, especially in high reward systems. We present empirical evidence that is consistent with this hypothesis. In terms of the consequences for plant fitness, we propose that specialized plant-pollinator systems should be more vulnerable to increased predation risk, with a steeper and faster decline in plant fitness, compared with generalist systems, in which pollinator redundancy can delay or buffer the effect of predators. Moreover, whereas we expect that specialist systems follows a similar inverse S-shape curve, in generalist systems we propose three different scenarios as a function not only of reward level but also compatibility, mating-system, and the interplay between growth form and floral display. The incorporation of trade-offs in pollinator behavior balancing the conflicting demands between feeding and predation risk has a promising future as a key feature enabling the development of more complex foraging models.
ABSTRACT
Although predators and floral herbivores can potentially decrease plant fitness by changing pollinator behaviors, studies comparing the strength of these factors as well as their additive and interactive effects on pollinator visitation and plant fitness have not been conducted. In this study, we manipulated the floral symmetry and predator presence (artificial crab spiders) on the flowers of the shrub Rubus rosifolius (Rosaceae) in a 2 × 2 factorial randomized block design. We found that asymmetry and predators decreased pollinator visitation (mainly hymenopterans), and overall these factors did not interact (additive effects). The effect of predation risk on pollinator avoidance behavior was 62 % higher than that of floral asymmetry. Furthermore, path analyses revealed that only predation risk cascaded down to plant fitness, and it significantly decreased fruit biomass by 33 % and seed number by 28 %. We also demonstrated that R. rosifolius fitness is indirectly affected by visiting and avoidance behaviors of pollinators. The strong avoidance behavioral response triggered by predation risk may be related to predator pressure upon flowers. Although floral asymmetry caused by herbivory can alter the quality of resources, it should not exert the same evolutionary pressure as that of predator-prey interactions. Our study highlights the importance of considering simultaneous forces, such as predation risk and floral asymmetry, as well as pollinator behavior when evaluating ecological processes involving mutualistic plant-pollinator systems.