Higher pollinator effectiveness by specialist than generalist flower-visitors of unspecialized Knautia arvensis (Dipsacaceae)

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Abstract
A critical issue in pollination ecology is the evolution of generalist pollination systems, and under which conditions floral specializations evolve from these. The gynodioecious herb Knautia arvensis (Dipsacaceae) exhibits a generalized pollination system, but is visited by both generalist and specialist flower-visitors. The present study tested pollinator effectiveness and pollinator importance of the pollen specialist solitary bee Andrena hattorfiana (Andrenidae) vs. the generalist flower-visitors to K. arvensis on the island of Öland, SE Sweden. Females of the specialist bee removed more pollen per inflorescence-visit than the major groups of generalist visitors such as bumblebees and flies. They also deposited more pollen per inflorescence-visit than any of the generalist visitor groups. The females have a preference for pollen-presenting vs. stigma-presenting inflorescences, a pattern shared with most of the generalist flower-visitors. Females of the specialist exert such a strong preference that they, despite their great pollinator effectiveness, make modest contribution to pollen transfer in K. arvensis. The females of A. hattorfiana accounted for 14.2% of the overall visits and 5.8% of the total pollination, the rest being performed by generalist visitors and males of A. hattorfiana. This study shows that pollinator effectiveness of a specialist can be superior while generalist flower-visitors select floral characters towards generalization through their greater contribution to overall pollen flow.