Competition for bumblebee visitation between Melampyrum pratense and Viscaria vulgaris with healthy and Ustilago-infected flowers

We studied bumblebee foraging on two sympatrically and simultaneously flowering species, Melampyrum pratense (Scrophulariaceae) and Viscaria vulgaris (Caryophyllaceae) during the flowering season of Viscaria in south-west Sweden. We distinguished between healthy and Ustilago-infected Viscaria plants. Both species shared the main insect visitor, queens of Bombus hortorum, which collected nectar from both species but pollen from Melampyrum only. The pattern of visitation changed over the season: bumblebees preferred Viscaria early on, but changed to Melampyrum later in the season, probably because of the higher sugar content of Melampyrum nectar and the possibility of collecting both nectar and pollen from the same flower. Pollen collecting is probably of increasing importance since the need of pollen for the developing larvae will increase with time. Flowers of Viscaria received fewer visits in plots with other species than in pure Viscaria plots during one year and received more visits early than late in the season during both years. Melampyrum flowers received similar amounts of visits in mixed and pure environments. They also received more visits early than late, although this was probably a result of pollinator satiation since Melampyrum became very abundant with time. Ustilago-infected plants received far fewer visits but because of its long flowering time the proportion of open flowers receiving visits was still high. Viscaria flowers received significantly more visits than flowers of other species when bumblebees made heterospecific flower visits from Ustilago-infected plants; thus Ustilago spores were probably effectively dispersed from infected to healthy plants by the pollinators. The mechanism behind competition for pollination in this system was competition through pollinator preference, since the visitation rate to Viscaria actually decreased, but also competition through improper pollen transfer (grains of both species were found on the bodies of bumblebees) since the incidence of switching between the two species increased, probably resulting in an increased misplacement of conspecific pollen grains with time.