In animal-pollinated plants, the opportunity for selection and the strength of pollinator-mediated selection are expected to increase with the degree of pollen limitation. However, whether differences in pollen limitation can explain variation in pollinator-mediated and net selection among animal-pollinated species is poorly understood. In the present study, we quantified pollen limitation, variance in relative fitness and pollinator-mediated selection on five traits important for pollinator attraction (flowering start, plant height, flower number, flower size) and pollination efficiency (spur length) in natural populations of 12 orchid species. Pollinator-mediated selection was quantified by subtracting estimates of selection gradients for plants receiving supplemental hand-pollination from estimates obtained for open-pollinated control plants. Mean pollen limitation ranged from zero to 0.96. Opportunity for selection, pollinator-mediated selection and net selection were all positively related to pollen limitation, whereas nonpollinator-mediated selection was not. Opportunity for selection varied five-fold, strength of pollinator-mediated selection varied three-fold and net selection varied 1.5-fold among species. Supplemental hand-pollination reduced both opportunity for selection and selection on floral traits. The results show that the intensity of biotic interactions is an important determinant of the selection regime, and indicate that the potential for pollinator-mediated selection and divergence in floral traits is particularly high in species that are strongly pollen-limited.
Contrasting flower color patterns that putatively attract or direct pollinators toward a reward are common among angiosperms. In the deceptive orchid Anacamptis morio, the lower petal, which makes up most of the floral display, has a light central patch with dark markings. Within populations, there is pronounced variation in petal brightness, patch size, amount of dark markings, and contrast between patch and petal margin. We tested whether pollinators mediate selection on these color traits and on morphology (plant height, number of flowers, corolla size, spur length), and whether selection is consistent with facilitated or negative frequency-dependent pollination. Pollinators mediated strong selection for increased petal brightness (Δβpoll = 0.42) and contrast (Δβpoll = 0.51). Pollinators also tended to mediate stabilizing selection on brightness (Δγpoll = -0.27, n.s.) favoring the most common phenotype in the population. Selection for reduced petal brightness among hand-pollinated plants indicated a fitness cost associated with brightness. The results demonstrate that flower color traits influence pollination success and seed production in A. morio, indicating that they affect attractiveness to pollinators, efficiency of pollen transfer, or both. The documented selection is consistent with facilitated pollination and selection for color convergence toward cooccurring rewarding species.
Most plants attract multiple flower visitors that may vary widely in their effectiveness as pollinators. Floral evolution is expected to reflect interactions with the most important pollinators, but few studies have quantified the contribution of different pollinators to current selection on floral traits. To compare selection mediated by diurnal and nocturnal pollinators on floral display and spur length in the rewarding orchid Gymnadenia conopsea, we manipulated the environment by conducting supplemental hand-pollinations and selective pollinator exclusions in two populations in central Norway. In both populations, the exclusion of diurnal pollinators significantly reduced seed production compared to open pollination, whereas the exclusion of nocturnal pollinators did not. There was significant selection on traits expected to influence pollinator attraction and pollination efficiency in both the diurnal and nocturnal pollination treatment. The relative strength of selection among plants exposed to diurnal and nocturnal visitors varied among traits and populations, but the direction of selection was consistent. The results suggest that diurnal pollinators are more important than nocturnal pollinators for seed production in the study populations, but that both categories contribute to selection on floral morphology. The study illustrates how experimental manipulations can link specific categories of pollinators to observed selection on floral traits, and thus improve our understanding of how species interactions shape patterns of selection.
Flower longevity is an adaptive trait, optimized to balance reproductive success against the costs of flower maintenance. The trait is highly plastic in response to pollination success, and numerous studies report increased flower longevity in high elevation environments, where diversity, abundance, and activity of pollinators are low. However, few studies have experimentally investigated how flower longevity varies with pollination intensity within and among populations. We studied flower longevity of six alpine species under three pollination intensity treatments (hand-pollination, natural pollination, pollinator exclusion) at 1600 m vs. 2600 m a.s.l. at the Furka Pass, Central Swiss Alps. We hypothesized, (1) that flower longevity is generally increased in population at high elevation, and (2) that the increase in flower longevity when pollination fails is stronger in populations at high elevation compared to low elevation. Hand-pollination did not decrease flower longevity in any of the studied populations and rarely increased natural seed production suggesting no pollination limitation at both elevations. This was supported by similar pollinator visitation rates, pollinator efficiency, and pollination effectivity. Pollinator exclusion significantly increased flower longevity, but only in populations of three species at low elevation, whereby in all populations of the six species at high elevation, indicating a higher plasticity of flowers in populations at high elevation compared to populations from lower elevation. We suggest that the higher plasticity of flower longevity in alpine populations is of advantage in their unpredictable pollination environment: Increased flower longevity compensates for low pollination in unsuitable periods guaranteeing a minimum reproduction, while the capacity to senescence rapidly after successful pollination saves redundant floral costs in suitable periods.
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