'X' marks the spot: The possible benefits of nectar guides to bees and plants

1. Many floral displays are visually complex, transmitting multi-coloured patterns that are thought to direct pollinators to nectar rewards. These nectar guides may be mutually beneficial, if they reduce pollinators handling time, leading to an increased visitation rate and promoting pollen transfer. Yet, many details regarding how floral patterns influence foraging efficiency are unknown, as is the potential for pollinator learning to alter this relationship. 2. We compared the responses of bumblebee (Bombus impatiens Cresson) foragers to artificial flowers that either possessed or lacked star-like patterns. By presenting each bee with two different foraging scenarios (patterned flowers rewarding/plain flowers unrewarding, plain flowers rewarding/patterned flowers unrewarding) on different days, we were able to assess both short- and long-term effects of patterns on bee foraging behaviour. 3. Bees discovered rewards more quickly on patterned flowers and were less likely to miss the reward, regardless of whether corollas were circular or had petals. Nectar guides' effect on nectar discovery was immediate (innate) and persisted even after experience, although nectar discovery itself also had a learned component. We also found that bees departed patterned flowers sooner after feeding. Finally, when conditions changed such that flowers no longer provided a reward, bees visited the now-unrewarding flowers more persistently when they were patterned. 4. On the time-scale of a single foraging bout, our results provide some of the first data on how pollinators learn to forage efficiently using this common floral trait. Our bees' persistent response to patterned flowers even after rewards ceased suggests that, rather than being consistently mutually beneficial to plant and pollinator, nectar guide patterns can at times promote pollen transfer for the plant at the expense of a bee's foraging success.