Adaptive pattern of nectar volume within inflorescences: bumblebee foraging behavior and pollinator-mediated natural selection

Larger floral displays increase pollinator visitation as well as among-flower self-pollination (geitonogamy) in self-compatible species. Dichogamy (temporal separation of gender expression) can limit geitonogamy and increase outcrossing but this depends on pollinator behavior within inflorescences. Declining nectar volume from lower to upper flowers is a hypothesized adaptation to increase outcrossing and pollen export by encouraging the upward movment of pollinators from female to male flowers and by reducing the number of flowers probed per inflorescence, but supporting evidence has been equivocal. We tested this hypothesis in Aconitum gymnandrum by studying floral display and rewards, pollinator visitation, and pollinator-mediated selection on floral traits. We found that larger inflorescences of A. gymnandrum attracted more pollinators, but did not increase the number of flowers probed per visit. Nectar production declined with increasing flower height on average, but the opposite pattern was also common. Bumblebees responded strongly to the nectar pattern, moving from higher to lower nectar concentration. Finally, there was significant pollinator-mediated direct selection for this pattern of declining nectar volume after correcting for correlations with flower size, number, and mean nectar volume. Together, the results strongly suggest that declining nectar production in higher flowers is an adaptation to enhance outcrossing in A. gymnandrum.