Pollination-precision hypothesis: support from native honey bees and nectar bats.

The evolution of floral traits is often considered to reflect selection for increased pollination efficiency. Known as the pollination-precision hypothesis, increased pollination efficiency is achieved by enhancing pollen deposition on precise areas of the pollinator. Most research to date addressing this hypothesis has examined plant species that are a priori predicted to place pollen precisely, but we still lack comparisons with species predicted to have low pollination efficiency. We studied 39 plant species with diverse floral morphologies and measured the precision of pollen placement on two pollinator groups: honey bees (genus Apis) and nectar bats (family Pteropodidae). Pollen was collected from four locations of each pollinator's body (bees: dorsal thorax, ventral thorax, dorsal abdomen, ventral abdomen; bats: crown, face, chest, wing) to calculate pollen placement precision using Pielou's evenness index. We also quantified variation in floral design by scoring floral symmetry, corolla fusion, floral orientation and stamen number. We confirm the importance of four floral character states (bilateral symmetry, fused corollas, horizontal orientation and reduced stamen number) in promoting precise pollen placement on diverse pollinators. Our findings provide phylogenetically corrected, empirical support that the evolution of the four floral characters reflect selection for enhanced precision of pollen placed on pollinators.

Comparison of population genetic structures of the plant Silene stellata and its obligate pollinating seed predator moth Hadena ectypa.

Background and Aims: Population genetic structures and patterns of gene flow of interacting species provide important insights into the spatial scale of their interactions and the potential for local co-adaptation. We analysed the genetic structures of the plant Silene stellata and the nocturnal moth Hadena ectypa. Hadena ectypa acts as one of the important pollinators of S. stellata as well as being an obligate seed parasite on the plant. Although H. ectypa provides a substantial pollination service to S. stellata, this system is largely considered parasitic due to the severe seed predation by the Hadena larvae. Previous research on this system has found variable interaction outcomes across space, indicating the potential for a geographical selection mosaic. Methods: Using 11 microsatellite markers for S. stellata and nine markers for H. ectypa, we analysed the population genetic structure and the patterns and intensity of gene flow within and among three local populations in the Appalachians. Key Results: We found no spatial genetic structure in the moth populations, while significant differentiation was detected among the local plant populations. Additionally, we observed that gene flow rates among H. ectypa populations were more uniform and that the mean gene flow rate in H. ectypa was twice as large as that in S. stellata. Conclusions: Our results suggest that although the moths move frequently among populations, long-distance pollen carryover only happens occasionally. The difference in gene flow rates between S. stellata and H. ectypa could prevent strict local co-adaptation. Furthermore, higher gene flow rates in H. ectypa could also increase resistance of the local S. stellata populations to the parasitic effect of H. ectypa and therefore help to stabilize the Silene-Hadena interaction dynamics.

Choices and consequences of oviposition by a pollinating seed predator, Hadena ectypa (Noctuidae), on its host plant, Silene stellata (Caryophyllaceae).

PREMISE OF THE STUDY: Pollinating seed predators are models for the study of mutualisms. These insects have dual effects on host-plant fitness, through pollination as adults and flower and fruit predation as larvae. A rarely examined question is whether pollinating seed-predator oviposition choices are influenced by plant floral and size traits and the potential consequences of oviposition for host-plant reproduction. • METHODS: We quantified oviposition by a pollinating seed predator, Hadena ectypa, on its host, Silene stellata, to determine if oviposition was associated with specific plant traits and whether oviposition was significantly correlated with fruit initiation or flower and fruit predation over three years. We also quantified whether stigmatic pollen loads of flowers visited by Hadena that both fed on nectar and oviposited were greater than when Hadena only fed on nectar. • KEY RESULTS: Hadena had significant preference for plants having flowers with long corolla tubes in all three years. Moth oviposition was correlated with other traits only in some years. Oviposition did not increase stigmatic pollen loads. We observed significant positive relationships between both oviposition and fruit initiation and oviposition and flower/fruit predation. • CONCLUSIONS: Hadena ectypa oviposition choices were based consistently on floral tube length differences among individuals, and the consequences of oviposition include both fruit initiation (due to pollination while feeding on nectar prior to oviposition) and larval flower/fruit predation. The positive association between oviposition and fruit initiation may explain the long-term maintenance of facultative pollinating seed-predator interactions.

Experimental floral and inflorescence trait manipulations affect pollinator preference and function in a hummingbird-pollinated plant.

PREMISE OF THE STUDY: Controversy is ongoing regarding the importance of pollinator-mediated selection as a source of observed patterns of floral diversity. Although increasing evidence exists of pollinator-mediated selection acting on female reproductive success, there is still limited understanding of pollinator-mediated selection on floral traits via male reproductive success. Here we quantify potential selection by the ruby-throated hummingbird, Archilochus colubris, on four floral traits of hermaphroditic Silene exerted through male floral function. METHODS: In single trait manipulative experiments we quantified hummingbird visitation preference and/or fluorescent dye (a pollen analog) donation as a function of number of flowers displayed (inflorescence size), height of the floral display (inflorescence height), floral color, and corolla tube length. KEY RESULTS: Hummingbirds preferred to visit larger floral displays and floral displays at greater height, likely representing a general pollinator preference for larger, more visible signals and/or greater rewards. In addition, hummingbirds preferred to visit red flowers, and male function was greater in flowers manipulated to have longer corolla tubes. CONCLUSIONS: Selection pressures exerted by hummingbirds on S. virginica floral and inflorescence design through male reproductive success are consistent with the contemporary expression of floral traits of S. virginica relative to related Silene species with different pollinators, and they are consistent with the hummingbird syndrome of traits expressed by S. virginica.

Pollen limitation and reproductive assurance in Antillean Gesnerieae: a specialists vs. generalist comparison.

Pollen limitation of female fecundity is widespread among angiosperms, a signal that pollinators frequently fail to transfer pollen to fertilize all ovules. Recent surveys have suggested that pollen limitation is associated with floral specialization. This study uses a group of Antillean Gesneriaceae with contrasting pollination systems (bat, hummingbird, and generalist) to assess the premise that plants with specialized pollination systems and infrequent pollinator visitation experience greater pollen limitation of fruit and seed set than their generalist congeners. Alternatively, specialists may possess mechanisms that reduce pollen limitation, such as autonomous self-pollination. A survey of autonomous self-pollination conducted on 13 Gesneria and Rhytidophyllum species during 2006-2008 revealed no significant association between reproductive assurance mechanisms and pollination system specialization. However, high levels of potential autonomous self-pollination were only found among specialized hummingbird-pollinated species. A comparison of fruit and seed set between emasculated and unmanipulated flowers provided evidence for autonomous selfing acting as a reproductive assurance mechanism in three out of four ornithophilous species. Furthermore, the Puerto Rican population of G. reticulata relies almost exclusively on self-pollination for reproduction. Two-year pollen supplementation experiments conducted on nine Gesnerieae species from the Dominican Republic and Puerto Rico provided evidence for significant pollen limitation associated with pollination specialization including both bat- and hummingbird-pollinated Gesnerieae; no pollen limitation was detected in any of the four generalist species. No pollen limitation was detected either in two ornithophilous Gesneria species with low hummingbird visitation and high levels of autonomous self-pollination. This study provides support for the idea that generalized pollination systems may, in some cases, buffer against fluctuations in the pollinator environment. However, the use of reproductive assurance mechanisms allows the maintenance of highly specialized pollination systems in pollinator depauperate environments.

Pollination and seed predation by moths on Silene and allied Caryophyllaceae: evaluating a model system to study the evolution of mutualisms.

Nursery pollinators, and the plants they use as hosts for offspring development, function as exemplary models of coevolutionary mutualism. The two pre-eminent examples--fig wasps and yucca moths--show little variation in the interaction: the primary pollinator is an obligate mutualist. By contrast, nursery pollination of certain Caryophyllaceae, including Silene spp., by two nocturnal moth genera, Hadena and Perizoma, ranges from antagonistic to potentially mutualistic, offering an opportunity to test hypotheses about the factors that promote or discourage the evolution of mutualism. Here, we review nursery pollination and host-plant interactions in over 30 caryophyllaceous plants, based on published studies and a survey of researchers investigating pollination, seed predation, and moth morphology and behavior. We detected little direct evidence of mutualism in these moth-plant interactions, but found traits and patterns in both that are nonetheless consistent with the evolution of mutualism and merit further attention.