Summary 1.Pollinator visitation patterns in relation to variation in floral display size may be modified both quantitatively and qualitatively by local plant density. In this study four measures of pollinator response by Bombus spp. (plant visitation rate, bout length, proportion of flowers visited, flower visitation rate) were investigated under two or three different plant densities in two consecutive years in a natural population of Digitalis purpurea L. 2. Plant visitation rate increased with floral display size in both years, and was higher in dense patches compared with sparse ones in 1999. Bout lengths increased with display size in 1999, and bouts were longer in sparse patches. However, the actual rate of increase with display size was independent of plant density for both response measures. 3. The proportion of flowers visited decreased with floral display size in both years, and in 1999 the decline was faster in high-density patches. As a result, the proportion visited was higher in dense patches for the smallest display sizes, and higher in sparse patches for larger display sizes. 4. Flower visitation rate decreased with floral display size in both years. This is inconsistent with the idea that bees achieve an ideal free distribution across flowers. There was no significant effect of plant density. 5. These results demonstrate that local plant density variation may modify the functional relationship between floral display size and pollinator visitation rate, and potentially influence plant mating patterns.

Summary• Nonrewarding animal-pollinated plants commonly experience severe pollen limitation, which should result in strong selection on traits affecting the success of pollination. However, the importance of pollinators as selective agents on floral traits in deceptive species has not been quantified experimentally.• Here, we quantified pollinator-mediated selection (Db poll ) on floral morphology and start of flowering in the deceptive orchid Dactylorhiza lapponica by subtracting estimates of selection gradients for plants receiving supplemental handpollination from estimates obtained for open-pollinated control plants.• There was directional selection for taller plants with more flowers and longer spurs, but no statistically significant selection on corolla size or flowering start. Pollinator-mediated selection accounted for all observed selection on spur length (Db poll = 0.32), 76% of the selection on plant height (Db poll = 0.19) and 42% of the selection on number of flowers (Db poll = 0.30). Sixteen per cent of developing fruits were consumed by insect herbivores, but fruit herbivory had only minor effects on the strength of pollinator-mediated selection.• Our results demonstrate that pollinators mediate selection on floral traits likely to affect both pollinator attraction and pollination efficiency, and are consistent with the hypothesis that deceptive species experience strong selection for increased display and mechanical fit between flower and pollinator.

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Abstract. Clarifying the relationship between environmental context and the adaptive significance of floral traits is fundamental for an understanding of spatial and temporal variation in pollinator-mediated selection. We manipulated vegetation height and pollination regime of the orchid Dactylorhiza lapponica in a factorial design to test whether pollinatormediated selection on floral traits is stronger in tall than in short vegetation, and whether this difference is larger for visual traits affecting pollinator attraction than for traits affecting pollination efficiency. In tall vegetation, pollinators mediated strong selection for taller plants (change in selection gradient for pollination, Db poll ¼ 0.33), more flowers (Db poll ¼ 0.34), and longer spurs (Db poll ¼ 0.42). In short vegetation, there was no significant selection on plant height, and pollinator-mediated selection on number of flowers and spur length was reduced by 52% and 25%, respectively. The results demonstrate experimentally that vegetation context can markedly influence the strength of pollinator-mediated selection on visual display traits, and indicate that this effect is weaker for traits affecting pollination efficiency. The study illustrates how crossed manipulations of environmental factors can reveal the causal links between ecological context and selection on floral traits.

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