Heterospecific pollen avoidance strategy prevails in the generalized plant-pollinator network on Yongxing Island.

Heterospecific pollen (HP) deposition varies widely among species in communities, which has been explicated by two adaptation strategies: HP avoidance and HP tolerance. Studies of the plant-pollinator network have uncovered that oceanic island communities are highly generalized and strongly connected. It remains unclear, however, which strategy prevails in such communities. We examined stigma pollen deposition on 29 plant species, and assessed patterns of HP load size and diversity in the Yongxing Island community. We assessed the effects of phenotypic specialization and species-level network structural properties of plant species on pollen deposition among species. The hypothesis of three accrual patterns of HP within species was tested by illustrating the relationship between conspecific pollen (CP) and HP receipt. Extensive variation occurred among species in HP receipt, while 75.9% of species received less than 10% HP and one species received more than 40% HP throughout the community. Flower size strongly drives the variation of HP receipt, while network structural properties had no effect on the pollen receipt. Nineteen species showed no relationship between the number of HP and CP loads, and they received smaller HP load sizes and lower HP proportions. Most plant species evolved HP avoidance strategy, and HP receipt was an occasional event for most plant species in the generalized community. HP and CP receipts are independent of each other in plant species with the HP avoidance mechanism. Our results highlight that plants in the generalized pollination system may preferentially select to minimize the HP load on stigmas.

Investigating the longstanding mystery of stigma length differences between morphs of distylous plants.

Longer stigmas in short-styled morphs of distylous plants have been considered an adaptive characteristic that increases intermorph pollen deposition. The greater pollen deposition in short-styled stigmas may be a by-product of their longer length, making deposition effectiveness comparison between morphs unfeasible. Thus, investigating which morph has the relatively most efficient stigma (i.e., pollen deposition per unit length) can boost our understanding of the adaptive significance of longer stigmas. Here, we compared pollen deposition between morphs relative to stigma length and assessed whether short-styled stigmas are more, less, or equally effective in receiving pollen grains per unit length. We reviewed the literature to characterize the extent of sigma length differences between morphs using the two most speciose distylous genera as model systems: Palicourea and Psychotria (Rubiaceae). Then, we conducted a between-morph comparison of raw and relative pollen depositions in a Palicourea rigida population. We confirmed that short-styled stigmas were longer than long-styled stigmas in both Palicourea (194.31% longer) and Psychotria (52.59% longer) flowers. Furthermore, in the focal Pal. rigida, although short-styled stigmas were 268.04% longer and received 97.04% more pollen grains than long-styled stigmas, the relative pollen deposition had a reverse pattern, with short-styled stigmas being two-times less efficient in receiving pollen. Our results indicate that the longer length of the short-styled stigmas may have a compensatory effect, increasing the chances of pollen grains reaching the stigma and probably maintaining disassortative intermorph pollination in distylous plants.

Reproductive isolation between two sympatric bat-pollinated Bauhinia (Leguminosae).

Several barriers contribute to reproductive isolation between plant species, which can be classified as pre- or post-pollination. Understanding the strength of these barriers could clarify the factors that maintain reproductive isolation and thus species integrity. In this study, we quantified reproductive isolation between two bat-pollinated co-occurring Bauhinia species (B. acuruana and B. pentandra) with similar flower morphology. Over the course of 18 months, we assessed reproductive isolation between these two Bauhinia species by quantifying the individual strengths and absolute contributions of five pre- and post- pollination barriers. Our data showed that both species are completely isolated in their reproduction by a combination of several barriers. Although they co-occur in a few populations, we found a high degree of geographic isolation between them. And although their flowering periods overlap, there is a significant difference in flowering peaks. Both species have the same pollinating bats, but the interspecific transfer of pollen between the plant species may be reduced due to the different length of the flower stamens, resulting in different pollen deposition on the bats' bodies. We have documented complete incompatibility between taxa and conclude that pre- and post-pollination barriers are important factors in preventing gene flow, even in contact zones between these two species of Bauhinia. We highlight that our work is the first study to use methods to estimate the strength of reproductive isolation barriers between bat-pollinated species.

Staminal hairs increase pollinator attraction and pollination accuracy in Tradescantia fluminensis (Commelinaceae).

Staminal hairs are the particular appendages of stamens, which may affect pollinator foraging behaviour and pollen transfer. However, experimental evidence of the functions of staminal hairs in pollination remains scarce. Here, we conducted staminal hair manipulation experiments in Tradescantia fluminensis (Commelinaceae) to investigate their effects on visitation and pollen transfer by bees. Our observations revealed that both visitation rates and visit duration of honeybees (Apis cerana) to control flowers were significantly higher than that of hairless flowers. Moreover, removing the staminal hairs significantly decreased pollen deposition by honeybees (A. cerana), but did not affect pollen removal. The staminal hair was similar in length to the stamen and the pistil of T. fluminensis. The staminal hairs provide more footholds for honeybees, and they lay prone on the staminal hairs to collect pollen, which increased the accuracy of pollination through the consistent pollen placement and pick-up on the ventral surface of honeybees. These results showed that the staminal hairs in T. fluminensis may represent an adaptation to attract pollinators and enhance pollination accuracy.

Pollen grading prediction scale for patients with Artemisia pollen allergy in China: A 3-day moving predictive model.

BACKGROUND: Artemisia pollen is the most prevalent outdoor aeroallergen causing respiratory allergies in Beijing, China. Pollen allergen concentrations have a direct impact on the quality of life of those suffering from allergies. Artemisia pollen deposition grading predictions can provide early warning for the protection and treatment of patients as well as provide a scientific basis for allergen specific clinical immunotherapy. OBJECTIVE: To develop a model of Artemisia pollen grading to predict development in patients with pollen allergy. METHODS: Artemisia pollen data from four pollen monitoring stations in Beijing as well as the number of Artemisia pollen allergen serum specific immunoglobulin E positive cases in Beijing Tongren Hospital from 2014 to 2016 were used to develop a statistical model of pollen deposition and provide optimised threshold values. RESULTS: A logarithmic correlation existed between the number of patients with Artemisia pollen allergy and Artemisia pollen deposition, and the average pollen deposition for three consecutive days was most correlated with the number of allergic patients. Based on the threshold of the number of patients and the characteristics of Artemisia pollen, a five-stage pollen deposition grading model was developed to predict the degree of pollen allergy. CONCLUSIONS: Graded prediction of pollen deposition may help pollen allergic populations benefit from preventive interventions before onset.

Stable pollination service in a generalist high Arctic community despite the warming climate.

Insects provide key pollination services in most terrestrial biomes, but this service depends on a multistep interaction between insect and plant. An insect needs to visit a flower, receive pollen from the anthers, move to another conspecific flower, and finally deposit the pollen on a receptive stigma. Each of these steps may be affected by climate change, and focusing on only one of them (e.g., flower visitation) may miss important signals of change in service provision. In this study, we combine data on visitation, pollen transport, and single-visit pollen deposition to estimate functional outcomes in the high Arctic plant-pollinator network of Zackenberg, Northeast Greenland, a model system for global warming-associated impacts in pollination services. Over two decades of rapid climate warming, we sampled the network repeatedly: in 1996, 1997, 2010, 2011, and 2016. Although the flowering plant and insect communities and their interactions varied substantially between years, as expected based on highly variable Arctic weather, there was no detectable directional change in either the structure of flower-visitor networks or estimated pollen deposition. For flower-visitor networks compiled over a single week, species phenologies caused major within-year variation in network structure despite consistency across years. Weekly networks for the middle of the flowering season emerged as especially important because most pollination service can be expected to be provided by these large, highly nested networks. Our findings suggest that pollination ecosystem service in the high Arctic is remarkably resilient. This resilience may reflect the plasticity of Arctic biota as an adaptation to extreme and unpredictable weather. However, most pollination service was contributed by relatively few fly taxa (Diptera: Spilogona sanctipauli and Drymeia segnis [Muscidae] and species of Rhamphomyia [Empididae]). If these key pollinators are negatively affected by climate change, network structure and the pollination service that depends on it would be seriously compromised.

Do Silene species with exposed stigmas tolerate interference by heterospecific pollen?

PREMISE: Co-flowering species that have not evolved an avoidance mechanism may have tolerance to heterospecific pollen (HP) deposition as an adaptive strategy to minimize any deleterious effects of HP transfer, but empirical evidence for the tolerance hypothesis remains scarce. METHODS: To estimate the potential effects of heterospecific pollen deposition (HPD) on female reproductive success, we counted conspecific (CP) and HP pollen grains deposited on stigmas and assessed subsequent seed set of both open- and hand-pollinated flowers in three co-flowering Silene species with exposed stigmas that usually received numerous HP grains on the elongated receptive area. RESULTS: The percentage of HP grains per flower (HP%) varied from 16.6% to 43.0% among three species. Silene chungtienensis had lower HP%, and the CP-HP relationship was neutral; S. gracilicaulis and S. yunnanensis had a relatively higher HP% with a positive CP-HP relationship. The effects of CP and HP number on natural seed set were positive for all three species, but HP% had stronger negative effects in S. chungtienensis and S. gracilicaulis. In hand-pollinated flowers of the three Silene species, seed set did not decrease with HP whether CP was in excess or insufficient, indicating no negative effects of HPD on seed production. CONCLUSIONS: Consistent with the tolerance hypothesis, our results indicated that species with higher HP interference are likely to be tolerant to an increase in HP%. These species with generalist-pollinated flowers and exposed large stigmas may benefit from an increase of conspecific pollen deposition, despite the associated increase in heterospecific pollen deposition.

Honey bee introductions displace native bees and decrease pollination of a native wildflower.

Introduced species can have cascading effects on ecological communities, but indirect effects of species introductions are rarely the focus of ecological studies. For example, managed honey bees (Apis mellifera) have been widely introduced outside their native range and are increasingly dominant floral visitors. Multiple studies have documented how honey bees impact native bee communities through floral resource competition, but few have quantified how these competitive interactions indirectly affect pollination and plant reproduction. Such indirect effects are hard to detect because honey bees are themselves pollinators and may directly impact pollination through their own floral visits. The potentially huge but poorly understood impacts that non-native honey bees have on native plant populations combined with increased pressure from beekeepers to place hives in U.S. National Parks and Forests makes exploring impacts of honey bee introductions on native plant pollination of pressing concern. In this study, we used experimental hive additions, field observations, as well as single-visit and multiple-visit pollination effectiveness trials across multiple years to untangle the direct and indirect impacts of increasing honey bee abundance on the pollination of an ecologically important wildflower, Camassia quamash. We found compelling evidence that honey bee introductions indirectly decrease pollination by reducing nectar and pollen availability and competitively excluding visits from more effective native bees. In contrast, the direct impact of honey bee visits on pollination was negligible, and, if anything, negative. Honey bees were ineffective pollinators, and increasing visit quantity could not compensate for inferior visit quality. Indeed, although the effect was not statistically significant, increased honey bee visits had a marginally negative impact on seed production. Thus, honey bee introductions may erode longstanding plant-pollinator mutualisms, with negative consequences for plant reproduction. Our study calls for a more thorough understanding of the indirect effects of species introductions and more careful coordination of hive placements.

Floral orientation affects outcross-pollen deposition in buzz-pollinated flowers with bilateral symmetry.

PREMISE: Floral orientation is central to plant-pollinator interactions and is commonly associated with floral symmetry. Bilaterally symmetrical flowers are often oriented horizontally for optimal pollinator positioning and pollen transfer efficiency, while the orientation of radially symmetrical flowers is variable. Buzz-pollinated species (pollinated by vibration-producing bees) include bilateral, horizontally oriented flowers, and radial, pendant flowers. The effect of floral orientation on pollen transfer has never been tested in buzz-pollinated species. METHODS: Here, we examined the effect of floral orientation on bumblebee-mediated pollen deposition in three buzz-pollinated Solanum species with different floral symmetry and natural orientations: S. lycopersicum and S. seaforthianum (radial, pendant), and S. rostratum (bilateral, horizontal). We tested whether orientation affects total stigmatic pollen deposition (both self and outcross pollen) when all flowers have the same orientation (either pendant or horizontal). In a second experiment, we evaluated whether different orientations of donor and recipient flowers affects the receipt of outcross pollen by S. rostratum. RESULTS: For the three Solanum species studied, there was no effect of floral orientation on total pollen deposition (both self and outcross) when flowers shared the same orientation. In contrast, in our experiment with S. rostratum, we found that pendant flowers received fewer outcross-pollen grains when paired with pendant donors. CONCLUSIONS: We suggest that floral orientation influences the quality of pollen transferred, with more outcross pollen transferred to horizontally oriented recipients in the bilaterally symmetrical S. rostratum. Whether other bilaterally symmetrical, buzz-pollinated flowers also benefit from increased cross-pollination when presented horizontally remains to be established.

Conspecific pollen advantage mediated by the extragynoecial compitum and its potential to resist interspecific reproductive interference between two Sagittaria species.

The extragynoecial compitum formed by the incomplete fusion of carpel margins, while allowing intercarpellary growth of pollen tubes in apocarpous angiosperms, may also increase the risk of reproductive interference caused by heterospecific pollen (HP) deposition. In Sagittaria, congeneric HP tubes grow via different paths and enter the ovules later than conspecific pollen (CP) tubes. However, it is unclear how the growth advantage of the CP tube helps ensure reproductive success when HP is deposited on the stigmas. We performed molecular characterization of interspecies-pollinated seeds to examine the consequences of interspecific pollen deposition between Sagittaria pygmaea and S. trifolia. We also conducted CP-HP (1:1) mixed pollination and delayed CP pollination treatments to explore the seed-siring abilities of CP and HP. Our results showed that although HP could trigger the development of fruits, the interspecies-pollinated seeds contained partially developed embryos and could not germinate. More than 70% of the embryos in these seeds were molecularly identified as hybrids of both species, suggesting that HP tubes could enter the ovules and fertilize the egg cells. Moreover, CP could sire more offspring (≥70%) after the CP-HP (1:1) mixed pollination treatment, even when HP reached the stigma 0.5-1 h earlier than CP (≥50%). Following adequate CP vs. HP (1:1) pollination on carpels on two sides of the apocarpous gynoecium, both species produced > 70% conspecific seeds, indicating that the CP tubes could occupy ovules that should be occupied by HP via the extragynoecial compitum. Our results reveal that in Sagittaria, pollen deposition from co-existing congeneric heterospecies leads to interspecific seed discounting. However, the CP advantage mediated by the extragynoecial compitum is an effective strategy to mitigate the effects of interspecific pollen deposition. This study improves our understanding of how apocarpous angiosperms with an extragynoecial compitum can maintain species stability and mitigate the negative reproductive interference effect from sympatrically distributed related species.

Structural anther mimics improve reproductive success through dishonest signaling that enhances both attraction and the morphological fit of pollinators with flowers.

Numerous studies have identified traits associated with anther mimicry; however, the processes underlying floral deception remain poorly documented for these structures. We studied the importance of pollinator attraction and mechanical fit of anther mimics in Tritonia laxifolia (Iridaceae) and their relative contributions to reproductive success. To determine anther mimics role in attraction, we offered bees' binary choices to flowers painted with UV-absorbent and UV-reflecting paints. We also conducted preference experiments between flowers with excised anther mimics and unmanipulated controls, from which mechanical fit was assessed by allowing single visits. Anther mimics' effects on female reproductive success were determined using similar treatments, but on rooted plants. Bees preferred UV-absorbent over UV-reflecting anther mimics. Bees did not discriminate between flowers with and without three-dimensional anther mimics. Single visits resulted in more pollen deposition on unmanipulated controls over flowers with their anther mimics excised, which was directly linked to pollen-collecting behavior. Controls with unmanipulated anther mimics had higher seed set than those with their anther mimics excised. This study provides insights into pollinator-mediated selection on deceptive floral signals and shows that three-dimensional anther mimics increases reproductive success through both attraction and pollen-collecting behaviors.

Hummingbird contribution to plant reproduction in the rupestrian grasslands is not defined by pollination syndrome.

Floral traits mediate the roles of distinct animals as effective pollinators along a generalization/specialization continuum. Many plant species are visited by different pollinator functional groups and the specific contribution of each group is expected to reflect the set of floral characteristics defined by pollination syndromes. Although considered a highly specialized nectarivorous group, hummingbirds frequently visit flowers lacking apparent specialization to bird pollination. How they contribute to the reproduction of these plants, however, has not been evaluated through field experiments considering multiple non-related plant species simultaneously. Here, we investigated hummingbirds' contributions to the pollination of ten plant species comprising a gradient of adaptation to bird pollination in the Brazilian rupestrian grasslands. We excluded hummingbirds from flowers and evaluated their relative contribution in comparison to insects (mainly bees) on conspecific/heterospecific pollen deposition and fruit set. Floral traits that are typically associated with bird pollination were associated with increased pollen deposition, but not with fruit set in the presence of hummingbirds. With hummingbirds, conspecific and heterospecific pollen deposition increased in most species, while fruit set increased in four plant species with varying degrees of fit to ornithophily. Our results show that assessing the relative contribution of specific pollinator groups may depend on when this contribution is measured, i.e. pollen deposition or fruit set. Considering fruit set, our results indicate that hummingbirds contributed to plant reproduction independently of the fit to bird pollination syndrome. This emphasizes their importance as under-appreciated generalized pollinators in some communities.

A meta-analysis of single visit pollination effectiveness comparing honeybees and other floral visitors.

PREMISE: Many animals provide ecosystem services in the form of pollination including honeybees, which have become globally dominant floral visitors. A rich literature documents considerable variation in single visit pollination effectiveness, but this literature has yet to be extensively synthesized to address whether honeybees are effective pollinators. METHODS: We conducted a hierarchical meta-analysis of 168 studies and extracted 1564 single visit effectiveness (SVE) measures for 240 plant species. We paired SVE data with visitation frequency data for 69 of these studies. We used these data to ask three questions: (1) Do honeybees (Apis mellifera) and other floral visitors differ in their SVE? (2) To what extent do plant and pollinator attributes predict differences in SVE between honeybees and other visitors? (3) Is there a correlation between visitation frequency and SVE? RESULTS: Honeybees were significantly less effective than the most effective non-honeybee pollinators but were as effective as the average pollinator. The type of pollinator moderated these effects. Honeybees were less effective compared to the most effective and average bird and bee pollinators but were as effective as other taxa. Visitation frequency and SVE were positively correlated, but this trend was largely driven by data from communities where honeybees were absent. CONCLUSIONS: Although high visitation frequencies make honeybees important pollinators, they were less effective than the average bee and rarely the most effective pollinator of the plants they visit. As such, honeybees may be imperfect substitutes for the loss of wild pollinators, and safeguarding pollination will benefit from conservation of non-honeybee taxa.

Pollination adaptations of group-by-group stamen movement in a meadow plant with temporal floral closure.

Floral sexual organ (stamen and pistil) movements are selective adaptations that have different functions in male-female reproduction and the evolution of flowering plants. However, the significance of stamen movements in the spatial-temporal function and separation of male and female organs has not been experimentally determined in species exhibiting floral temporal closure. The current study investigated the role of slow stamen (group-by-group) movement in male-female sexual function, and the effect of stamen movement on pollen removal, male-male and male-female interference, and mating patterns of Geranium pratense, a plant with temporal floral closure. This species uses stamen group-by-group movement and therefore anther-stigma spatial-temporal separation. Spatial separation (two whorls of stamen and pistil length) was shown to be stronger than temporal separation. We found that stamen movements to the center of the flower increase pollen removal, and the most common pollinators visited more frequently and for longer durations during the male floral stage than during the female floral stage. Petal movements increased both self-pollen deposition rate and sexual interference in G. pratense. The fruit and seed set of naturally and outcrossed pollinated flowers were more prolific than those of self-pollinated flowers. Group-by-group stamen movement, dehiscence of stamens, pistil movement, and male-female spatial-temporal functional separation of G. pratense before floral temporal closure may prevent male-female and stamen-stamen interference and pollen discounting, and may increase pollen removal and cross-pollination.

Inter-morph pollen flow and reproductive success in a self-compatible species with stigma-height dimorphism: the influence of herkogamy and reciprocity.

Inter-morph pollen transfer and its dependence on herkogamy and reciprocity are not completely understood in species with stigma-height dimorphism. We asked whether total stigmatic pollen loads, inter-morph fraction of pollen load and reproductive success differed between morphs in Jasminum malabaricum, a species exhibiting stigma-height dimorphism. We tested whether higher herkogamy and reciprocity resulted in higher inter-morph pollen deposition and reproductive success. We quantified individual-level estimates of herkogamy, reciprocity, total stigmatic pollen load, inter-morph stigmatic pollen fraction and fruit set for both morphs in naturally occurring populations of J. malabaricum. Total pollen load was higher in the long-styled morph, inter-morph pollen fraction was higher in the short-styled morph, but fruit set did not differ between morphs. Higher herkogamy resulted in a higher inter-morph fraction of pollen load and fruit set in the long-styled morph of one population. In the other population, only reciprocity was found to be related to inter-morph pollen deposition. This study is the first to quantify and report natural inter-morph stigmatic pollen load in a species with stigma-height dimorphism. Morph-specific differences in pollen load were similar to patterns commonly observed in heterostylous species. The results highlight the importance of both herkogamy and reciprocity in facilitating inter-morph pollen transfer. Population-specific patterns indicate that local environmental factors determine the relative functional importance of herkogamy and reciprocity.

Water availability affects the relationship between pollen intensity and seed production.

Seed production can be affected by water availability and also depend on the amount (pollen intensity) and quality of pollen deposited. The way pollen receipt on the stigma translates into seeds produced follows that of a saturating dose-response. Not only can water availability and pollen intensity each influence seed production, these factors could interact in their effects on seed production. Changes to the relationship between seed production and pollen intensity can in turn influence pollinator effectiveness and pollinator-mediated selection. We asked how water availability affected indices of plant fitness (seed set, fruit set and seed mass) and the relationship between pollen intensity and seed production in Phacelia parryi. We conducted a greenhouse experiment where we manipulated water availability (either high- or low-water) to pollen recipient plants and hand-pollinated each plant with a range of pollen intensities. We conducted 703 hand-pollinations on 30 plants. For each hand-pollinated flower we measured pollen deposited, seed production and seed mass. We then generated a piecewise regression of the relationship between pollen intensity and seed production, and determined average effects of water on plant fitness measures. This experiment was paired with a field observational study aimed to document natural variation in pollen deposition. Average seed production per fruit was 21 % higher in the high-watered plants. The relationship between pollen intensity and seed production differed between the two water treatments. Plants under high-water exhibited a wider range in which pollen deposition increased seed production. Average natural pollen intensities fell within different regions of the piecewise regression for low- and high-water plants. Water availability can alter the efficiency by which pollen received is translated into seeds produced. Our greenhouse data suggest that only under certain pollen intensity environments will water availability affect how pollen received is translated into seeds produced.

Plant-pollinator network structural properties differentially affect pollen transfer dynamics and pollination success.

Plant-pollinator network studies have uncovered important generalities in the structure of these communities, rapidly advancing our understanding of the underlying drivers of such a structure. In spite of this, however, it is still unclear how changes in structural network properties influence overall plant pollination success. One key limitation is the lack of information on the relationship between network structural properties and aspects of pollination and plant reproductive success. Here, we estimate four plant species network structural metrics (interaction strength, weighted degree, closeness centrality, and specialization level), commonly used to describe their importance within plant-pollinator networks, at two different sites, and evaluate their effects on pollen deposition and pollen tube success. We found a positive effect of plant-pollinator specialization and a negative effect of closeness centrality on heterospecific pollen load size. We also found a marginal negative effect of closeness centrality on pollen tube success. Our results suggest that increasing plant-pollinator specialization within nested communities (pollinated by one or very few generalist insect species) may result in high levels of heterospecific pollen transfer. Furthermore, the differential effects of plant-pollinator network metrics on pollination success (pollen receipt and pollen tube success), highlight the need to integrate quantity (e.g. visitation rate) and quality (e.g. pollen delivery) aspects of pollination to achieve a more mechanistic understanding of the relationship between plant-pollinator network structure and function. Such knowledge is key to evaluate the resilience and stability of plant-pollinator communities and the services they provide in the face of increasing human disturbances.

The role of lateral and vertical herkogamy in the divergence of the blue- and red-flowered lineages of Lysimachia arvensis.

BACKGROUND AND AIMS: Herkogamy, or anther-stigma separation, is known to reduce self-pollen deposition, but little is known about the relative efficacy of different modes or conformations of herkogamy. We assessed the effectiveness of vertical versus lateral herkogamy in preventing or promoting self-pollen deposition in the annual herb Lysimachia arvensis, a plant with lineages that differ in flower colour, and in which flowers first display lateral and then vertical herkogamy. Because mating between the two lineages compromises fitness through the production of low-quality hybrid offspring, we tested the prediction that individuals sampled from sites occupied by both lineages should have flowers that promote autonomous self-pollen deposition and self-fertilization as a result of selection to reduce deleterious reproductive interference. METHODS: We characterized variation in herkogamy within and among 25 pure and mixed populations of L. arvensis in its European range and assessed the effectiveness of lateral versus vertical herkogamy in avoiding self-pollen deposition. RESULTS: Lateral herkogamy was more effective than vertical herkogamy in limiting self-pollen deposition. In the case of vertical herkogamy, only approach herkogamy was effective. Lineages showed consistent differences in herkogamy traits. In general, angles were smaller for blue than red flowers in most populations, and blue flowers showed approach herkogamy, while red flowers showed predominantly reverse herkogamy. In sympatry, the red lineage showed a reduction of both herkogamy traits while for the blue lineage only lateral herkogamy was reduced. CONCLUSIONS: Our results demonstrate that pollen deposition is affected not only by the degree but also the spatial conformation of herkogamy. They also highlight reduced herkogamy as a potential mechanism for promoting reproductive assurance under pollen limitation, as well as for avoiding reproductive interference between genetically divergent lineages.

Daily fluctuations in pollination effectiveness explain higher efficiency of native over exotic bees in Lepechinia floribunda (Lamiaceae).

BACKGROUND: Despite Stebbins' principle of the most efficient pollinator being proposed decades ago, the most important pollinators are still mainly identified using the frequency of visits to flowers. This shortcoming results in a gap between the characterization of the flower visitors of a plant species and a reliable estimation of the plant fitness consequences of the mutualistic interaction. The performance of a mutualistic visitor depends on its abundance, behaviour, effectiveness (pollen removal and deposition per unit time) and efficiency (seed set per unit time) conditioned by the temporal matching between pollinator activity and temporal patterns of maturation of the sexual functions of flowers. Although there have been recent attempts to provide a conceptual and methodological framework to characterize pollinators' performance, few have combined all key elements of visitors and plants to provide an accurate estimation of pollinators' performance under natural conditions. METHODS: We complement information on the flower biology and mating system of the sub-shrub Lepechinia floribunda (Lamiaceae) to provide a daily quantitative estimation of performance (effectiveness and efficiency) of the more abundant pollinators, i.e. native bumble-bees (Bombus spp.) and leafcutter bees (Megachile sp.), and the exotic honey-bee (Apis mellifera). KEY RESULTS: Unlike honey-bees or leafcutter bees, native bumble-bees matched the daily pattern of nectar production and stigma receptivity, and showed higher effectiveness and efficiency. Despite the overabundance of honey-bees, visits occurred mainly when stigmas were not receptive, thus reducing the honey-bees' overall performance. CONCLUSIONS: Bumble-bees appear to be the most important pollinators and potential historical mediators of reproductive trait evolution in L. floribunda. Because the production of seeds by bumble-bees involved fewer pollen grains for plants and less investment in floral display than honey-bees, contemporary and expected changes in pollinator abundance may affect future L. floribunda floral evolution. If bumble-bees were to be further displaced by anthropogenic disturbance or by competition with honey-bees, their lower efficiency will select for a larger floral display increasing reproductive costs. This scenario may also impose selection to reduce dichogamy to match honey-bee foraging activity.

Relative effectiveness of insects versus hummingbirds as pollinators of Rubiaceae plants across elevation in Dominica, Caribbean.

Most angiosperms rely on animal pollination for reproduction, but the dependence on specific pollinator groups varies greatly between species and localities. Notably, such dependence may be influenced by both floral traits and environmental conditions. Despite its importance, their joint contribution has rarely been studied at the assemblage level. At two elevations on the Caribbean island of Dominica, we measured the floral traits and the relative contributions of insects versus hummingbirds as pollinators of plants in the Rubiaceae family. Pollinator importance was measured as visitation rate (VR) and single visit pollen deposition (SVD), which were combined to assess overall pollinator effectiveness (PE). In the wet and cool Dominican highland, we found that hummingbirds were relatively more frequent and effective pollinators than insects, whereas insects and hummingbirds were equally frequent and effective pollinators at the warmer and less rainy midelevation. Furthermore, floral traits correlated independently of environment with the relative importance of pollinators, hummingbirds being more important in plant species having flowers with long and wide corollas producing higher volumes of dilute nectar. Our findings show that both environmental conditions and floral traits influence whether insects or hummingbirds are the most important pollinators of plants in the Rubiaceae family, highlighting the complexity of plant-pollinator systems.