During the evolution of angiosperm flowers, some floral traits may undergo certain changes in order to participate in screening. The stamens and pistils of Delphinium caeruleum are covered by two “door-like” staminodes, the evolutionary function of which, however, is quite unknown. In this study, we investigated whether D. caeruleum staminodes acted as visitor filters by assessing the respective strengths of staminodes and visitor insects (six bee species). We measured the operative strength required to open the staminodes and the strength that insects were capable of exerting using a biological tension sensor. Furthermore, we compared the strength required to open staminodes at different phases of the flowering period (male and female phases) and the strength of different visitors (visitors and non-visitors of D. caeruleum). The results showed that the strength needed to open staminodes in the male phase was significantly higher than that in the female phase. There was no significant difference between the strength exerted by visitors and required by staminodes of D. caeruleum in the male phase, but the visitor strength was significantly higher than that required to open staminodes in the female phase flowers. The strength of non-visitors was significantly lower than that required to open staminodes in the male phase. Furthermore, there was a significant positive association between the strength and the body weight of the bees. These results highlighted the observation that only strong visitors could press the two staminodes to access the sex organs and achieve successful pollination. Furthermore, these results revealed the function of pollinator screening by the staminodes of D. caeruleum. The biomechanical approach to the study of flowers allowed us to address relevant ecological and evolutionary questions of the plant–pollinator interaction and explore the functional modules within the flower structure in other plant species.
Nectar robbers, which affect plant fitness (directly or indirectly) in different degrees and in different ways, potentially constitute a significant part of mutualistic relationships. While the negative effects of nectar robbing on plant reproductive success have been widely reported, the positive effects remain unknown. The target of our study was to evaluate the effects of nectar robbers on the reproductive success of Symphytum officinale (Boraginaceae). We observed the behavior, species and times of visitors in the field, and we assessed the effect of nectar robbers on corolla abscission rate and time. To test the fitness of corolla abscission, we detected the changes in stigma receptivity, pollen viability, pollen amount and appendage opening size along with the time of flower blossom. The flowering dynamics and floral structure were observed to reveal the mechanism of self-pollination. Finally, pollen deposition seed set rate and fruit set rate were determined to estimate the effect of nectar robbers on reproduction success. We observed 14 species of visitors and 2539 visits in 50 h of observation; 91.7% of them were nectar robbers. The pressure and nectar removal of nectar robbers significantly promoted corolla abscission during a period when pollen grains are viable and the stigma is receptive. In addition, corolla abscission significantly increased the pollen deposition and seed setting rate. Our results demonstrate that nectar robbing contributes to enhancing seed production and positively and indirectly impacts the reproductive success of S. officinale. This mechanism involved the movement of anthers and indirect participation by nectar robbers, which was rarely investigated. Considering the multiple consequences of nectar robbing, understanding the impact of nectar robbers on plant reproduction is essential to comprehend the evolutionary importance of relationships between plants and their visitors.