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Pollination precision and efficiency have been deemed to be important driving forces in floral evolution. Herkogamy reduction is a main mechanism to increase pollination precision. Secondary pollen presentation (SPP), by which pollen is presented on other floral organs especially pistils, has been widely accepted as a special mechanism to increase pollen transfer precision through spatial reduction of the anther–stigma distance, that is, minimized herkogamy. This overlooks a potential driving force, that is expanding the pollination niche through converting pollen thieves and nectar robbers into effective pollinators. We selected two species as study models with typical pistillate SPP, Pavetta hongkongensis Bremek. (Rubiaceae) and Scaevola taccada (Gaertn.) Roxb. (Goodeniaceae). In both species, two distinct pollinator functional groups were recognized. Short‐tongued bees and flies fed on pollen on stigmas but also stole pollen from anthers and robbed nectar, whereas long‐tongued hawkmoths and butterflies only collected nectar. Emasculation had no influence on long‐tongued pollinators, but significantly decreased the visitation frequency of short‐tongued visitors and fruit set, compared to intact flowers, demonstrating short‐tongued visitors did not effectively pollinate and acted merely as pollen thieves or nectar robbers when SPP was absent. Data from the two plant species clearly indicated pistillate SPP has additional adaptive advantages of converting ineffective visitors into pollinators and consequently widening the pollination niche, which could help plants overcome environmental stochasticity. Our results suggest that multiple selective forces drive the evolution of SPP and the minimization of herkogamy. [ABSTRACT FROM AUTHOR]

The interactions between plants and their pollinators are the result of convergent evolution of floral attributes reflecting pressure exerted by pollinators. Nonetheless, the strategies employed by floral visitors to collect floral resources are extremely complex, and commonly involve theft or robbery in addition to pollination. We describe here the behavioral repertory of Apis mellifera during the collection of the floral resources, and evaluated the robbing rates of A. mellifera on the buds and flowers of Pyrostegia venusta during periods of intense and sparse flowering. We recorded the behaviors exhibited by foraging bees while collecting floral resources, quantified the numbers of floral buds and flowers with perforations in their corolla tissues, and determined whether that damage reduced nectar production. The evaluations were conducted during two distinct periods: during the period of intense flowering of P. venusta, and during the period of sparse flowering. Nectar robbing was observed during 93.4% of the visits of foraging A. mellifera bees, while nectar theft was observed during only 0.7% of the visits, and pollen theft during 5.9%. The robbing of floral buds and flowers was most intense during the period of heavy flowering. Flowers that had been intensely robbed secreted significantly less nectar than those non-robbed. The unusual nectar robbing activities of A. mellifera, especially during the period of intense flowering indicates an optimization of access to larger volumes of food resources. Our results therefore point to a major limitation of nectar per floral unit during the intense flowering period of P. venusta due to the high activity of nectar robbing by A. mellifera bees. [ABSTRACT FROM AUTHOR]

Keywords: Asian honey bees; Gnetum; gymnosperms; moths; nocturnal pollination; plant-insect interactions; pollen theft; tropical rainforests EN Asian honey bees Gnetum gymnosperms moths nocturnal pollination plant-insect interactions pollen theft tropical rainforests 1 4 4 12/03/21 20211201 NES 211201 Bees belong to modern groups of pollinators, which originated concurrently with new angiosperms (flowering plants), and might have contributed to the global gymnosperm-to-angiosperm turnover in the mid-Cretaceous (125-90 million years ago [Ma]; Cardinal and Danforth 2013, Peris et al. 2017). Our data suggest that diurnal honey bees disrupt the pollination of an insect-pollinated gymnosperm by reducing the pool of pollen available for nocturnal pollinators, resulting in decreased pollinator pollen loads and reduced seed set. Then, in bee-present season, we quantified honey bee visitation rate, pollen loads on honey bees and pollen remaining in anthers after honey bee visits to assess the intensity of theft. [Extracted from the article]

Background and Aims Most angiosperms rely on pollinators to transport pollen and effect fertilization. While some floral visitors are effective pollinators, others act as thieves, consuming pollen but effecting little pollination in return. The importance of pollen theft in male and female reproductive success has received little attention. Here, we examined if pollen consumption by flies altered pollen receipt and exacerbated pollen limitation for a bumblebee-pollinated plant, Polemonium foliosissimum (Polemoniaceae). Methods To examine the effect of pollen-thieving flies, we took a three-pronged approach. First, we used single-visit observations to quantify pollen removal and pollen deposition by flies and bumblebees. Second, we manipulated pollen in the neighbourhood around focal plants in two years to test whether pollen reduction reduced pollen receipt. Third, we combined pollen reduction with hand-pollination to test whether pollen thieving exacerbated pollen limitation. Polemonium foliosissimum is gynodioecious in most populations in the Elk Mountains of central Colorado, USA. Thus, we also tested whether pollen theft affected hermaphrodites and females differently. Results Flies removed significantly more pollen and deposited less pollen per visit than did bumblebees. Reduction of pollen in the neighbourhood around focal plants reduced pollen receipt in both years but only nearly significantly so in 2015. In 2016, plants were significantly pollen-limited; hand-pollination significantly increased seeds per fruit for both hermaphrodites and females. However, the reduction of pollen around focal plants did not exacerbate pollen limitation for either hermaphrodites or females. Conclusions Our results suggest that plants tolerate significant consumption of pollen by thieves and pollinators by producing ample pollen to feed both and fertilize available ovules. Our results demonstrate that pollen limitation in P. foliosissimum is driven by lack of effective pollinators rather than lack of pollen. Teasing out these effects elucidates the relative importance of drivers of reproductive success and thus the expected response to selection by different floral visitors. [ABSTRACT FROM AUTHOR]

Hybrid cauliflower production predominately relies on pollen transfer from hermaphrodite to female lines by honeybees. However, the presence of other pollinators may impact pollination success. Here, we investigate how honeybee visitation frequency and behaviour vary with plant sex and presence of blowflies and affect seed and pod set. We found substantial pollen limitation when honeybees were alone. This was likely due to their higher visitation to hermaphrodite flowers, infrequent transition from hermaphrodite to female flowers and high nectar theft in female flowers. Pollen foragers fed on nectar on hermaphrodite, but not female flowers. Moreover, when blowflies were present, the seed set was lower than that with honeybees alone. Our study highlights the importance of understanding the plant mating system and pollinator foraging behaviour with and without other species present in order to maximize seed set in hybrid crops. [ABSTRACT FROM AUTHOR]

Super-generalism is a pollinator trait where species obtain floral resources from a very wide range of plant species. Theoretical and empirical studies suggest that on islands with low pollinator diversity, such pollinators should evolve to exploit a very wide range of floral morphologies. Super-generalism has implications not only for securing pollination network stability, but also for the invasibility of potential weeds that require specialist pollinators in their original ranges. Here we expand earlier studies on bees in Fiji to include a wider range of bee-plant interactions for bees that have been recently introduced into Fiji as well as the endemic Fijian halictine bee, Homalictus fijiensis. Our data show that the endemic Fijian bee has a much wider range of floral hosts than introduced bees, and this extends to pollen larceny of solanoid plant species that are usually buzz pollinated. Importantly, solanoid plants were not visited by introduced bee species, including the honeybee Apis mellifera, which is usually regarded as a super-generalist. Our findings are important because they add critical support to the hypothesis that super-generalism evolves in insular ecosystems with low pollinator diversity and that this may make such ecosystems vulnerable to invasion by exotic weeds. However, insular super-generalists may also have potential to stabilize plant-pollinator networks and may also be effective pollinators for exotic crop species, and this needs to be further explored in agricultural settings. [ABSTRACT FROM AUTHOR]

(1) Background: Cardiocrinum cordatum (Thunb.) Makino (Liliaceae) is a forest perennial herb distributed in East Asia. Although flower visitors for this plant species have been well reported, their contribution to pollination remains unknown. (2) Methods: We evaluated pollination contribution for visitors of C. cordatum flowers in a natural cool temperate forest. We investigated visiting frequency, the number of pollen grains per body surface, fruit set, and the mean number of seeds per fruit produced after a single visit of each visiting species. Combining the results of these experiments, we determined the most important pollinators of this species. (3) Results: For the population investigated in the study, the three most essential pollinators were the bumblebee (Bombus diversus tersatus) (Apidae), sweat bee (Halictidae sp.), and marmalade hoverfly (Episyrphus balteatus) (Syrphidae). Additionally, we found that the contribution of a flower-visiting ant species (Myrmica ruginodis Nylander (s.l.)) (Formicidae) is small. (4) Conclusions: Pollinator contributions differed among flower visitors. Our results underscore the insufficiency of current information about flower-visiting species to evaluate pollination contribution.

Summary: Visual signals attractive to friends may also attract enemies.  The bright colors of anthers and pollen have generally been thought to attract pollinators. We hypothesize that visual crypsis of anthers, and particularly pollen, should be favored in flowering plants because protection from pollen collectors reduces the loss of male gametes.To understand adaptive strategies relating to the color of pollen, we measured the color of pollen, undehisced anther sacs, and their background, the corolla, with a spectrometer for 104 insect‐pollinated flowering species from a natural community in Hengduan Mountains, southwest China.The colors of anthers, pollen and corollas were diverse in these species. The color diversity of exposed pollen was significantly higher than that of concealed pollen (i.e. where anthers are enclosed or shielded by corollas). The color contrast between pollen and corolla was significantly smaller in species with exposed pollen than in those with concealed pollen. Unlike anther color, exposed pollen color tended to match its background corolla color.Our phylogenetic comparative analysis showed contrasting effects of pollen color patterns between flowers with exposed pollen and those with concealed pollen, revealing a strategy of hiding pollen from pollen thieves via visual crypsis. [ABSTRACT FROM AUTHOR]

More than 20,000 angiosperm species possess non-dehiscent anthers that open through small pores at the anther's tip. These flowers are visited by bees that use vibrations to remove pollen, a phenomenon known as buzz pollination. However, some floral visitors fail to transfer pollen efficiently, either through a mismatch of flower and insect size, or because they are unable to buzz-pollinate. These visitors collect pollen, but provide little or no pollination, behaving as pollen thieves. Although pollen theft is widespread in plants, few studies have quantified the incidence of pollen thieves for buzz-pollinated plants. We use observations of natural populations and floral manipulations of Solanum rostratum (Solanaceae) to investigate the incidence of pollen theft, find morphological and behavioural differences between pollinators and thieves, measure the pollination efficiency of visitors, and characterize the reproductive ecology of this herb. We found that most visitors act as thieves, with <20 % of all bees contacting the stigma. Insect visitors that regularly failed to contact the stigma (illegitimate visitors), included buzzing and non-buzzing bees, were significantly smaller, visited fewer flowers per bout, and stayed longer in each flower than (legitimate) visitors that regularly contact the stigma. Few flowers visited solely by illegitimate visitors set fruit. Our results show that S. rostratum requires insect visitation to set seed and natural populations experience moderate pollen limitation. We conclude that insect size, relative to the flower, is the main determinant of whether a visitor acts as a pollinator or a pollen thief in S. rostratum. [ABSTRACT FROM AUTHOR]

Background and Aims Pollen-collecting bees are among the most important pollinators globally, but are also the most common pollen thieves and can significantly reduce plant reproduction. The pollination efficiency of pollen collectors depends on the frequency of their visits to female(-phase) flowers, contact with stigmas and deposition of pollen of sufficient quantity and quality to fertilize ovules. Here we investigate the relative importance of these components, and the hypothesis that floral and inflorescence characteristics mediate the pollination role of pollen collection by bees. Methods For ten Aloe species that differ extensively in floral and inflorescence traits, we experimentally excluded potential bird pollinators to quantify the contributions of insect visitors to pollen removal, pollen deposition and seed production. We measured corolla width and depth to determine nectar accessibility, and the phenology of anther dehiscence and stigma receptivity to quantify herkogamy and dichogamy. Further, we compiled all published bird-exclusion studies of aloes, and compared insect pollination success with floral morphology. Key Results Species varied from exclusively insect pollinated, to exclusively bird pollinated but subject to extensive pollen theft by insects. Nectar inaccessibility and strong dichogamy inhibited pollination by pollen-collecting bees by discouraging visits to female-phase (i.e. pollenless) flowers. For species with large inflorescences of pollen-rich flowers, pollen collectors successfully deposited pollen, but of such low quality (probably self-pollen) that they made almost no contribution to seed set. Indeed, considering all published bird-exclusion studies (17 species in total), insect pollination efficiency varied significantly with floral shape. Conclusions Species-specific floral and inflorescence characteristics, especially nectar accessibility and dichogamy, control the efficiency of pollen-collecting bees as pollinators of aloes. [ABSTRACT FROM AUTHOR]

Pollen is unique among floral rewards in functioning as both a carrier of gametes and an attractant and nutritious resource for floral visitors. Animals that collect pollen without pollinating (pollen thieves) could reduce siring success of thieved plants and cause pollen limitation of seed set at the population level; however, such impacts on plant reproduction have not been demonstrated experimentally. To test these effects we added hives of native honey bees (Apis mellifera scutellata) to populations of a primarily bird-pollinated plant, Aloe maculata, in eastern South Africa. In field and aviary trials, bee addition increased pollen removal from anthers but decreased pollen deposition on stigmas, and so reduced both male and female pollination components. Further, total seed production decreased with hive addition in the aviary experiment and in three of four field populations, indicating that population-level pollen theft can also compromise reproductive success. In the field, naturally occurring allodapine bees also seemed to act as pollen thieves, outweighing the effect of honey bee hive addition at one of the four aloe populations. Our results highlight the importance of social bees as pollen thieves, even of plants that have evolved in their presence, and the role of dichogamy in promoting pollen theft. Given the commonness of both social bees and dichogamy, pollen theft is likely a much more common influence on floral ecology and evolution than suggested by the sparse literature. [ABSTRACT FROM AUTHOR]

Bees are frequent visitors to plants that, on the basis of floral traits, would be expected to be specialized for bird pollination. We conducted bird exclusion experiments to establish the contribution that bees make to the female component of pollination success of five "ornithophilous" Aloe species in the succulent-rich subtropical thicket vegetation in southeastern South Africa. Experiments showed that all five aloes are genetically self-incompatible and require cross-pollination for seed set. Seed set in flowers exposed to both bird and bee visitors was generally several fold higher than in flowers exposed to bee visitors only. The magnitude of this effect varied greatly among species and was affected by floral traits and bee behavior. In three aloes where the relatively short corolla tubes are blocked by exserted filaments, bees made a negligible contribution to seed production. However, in two aloes with long tubular flowers and included filaments, bees often crawled into the perianth tube to obtain nectar and contributed to seed production. Pollen-collecting honeybees contribute the least to pollination and may even impact negatively on aloe reproductive success by acting as pollen thieves. [ABSTRACT FROM AUTHOR]

Visual floral guides such as colored anthers, lines, dots, and UV-absorption patterns on petals are commonly observed in insect-pollinated angiosperms. Floral guides that are known to enhance foraging efficiency of visitors on flowers thus promote return visits (foraging facilitation hypothesis, which predicts that visitors will discriminate against flowers with inferior floral guides). In this study, we experimentally examined the hypothesis that floral guides also prevent pollen-theft behavior by foral visitors (theft prevention hypothesis), which has rarely been tested. Nectarless flowers of Commelina communis have three types of brightly colored floral organs: large blue petals, rewarding yellow anthers, and nonrewarding yellow anthers. Colored floral organs were removed artificially from plants in two natural populations of C. communis. Removal of the nonrewarding yellow anthers diminished hoverings in front of flowers and tended to reduce the number of total floral visitor landings, supporting the foraging facilitation hypothesis. Additionally, removal of the rewarding yellow anther decreased the frequency of legitimate landings on flowers and the legitimate landing-to-total landing ratio, which is consistent with the theft prevention hypothesis. The nonrewarding anthers and the rewarding yellow anthers were shown to play an important role in increasing visitor landings and orienting floral visitors toward a landing point appropriate for pollination, respectively. We also showed that the absence of yellow anthers decreased both pollen dispatch from brown anthers and receipt by stigmas in C. communis. These findings support both the foraging facilitation hypothesis and the theft prevention hypothesis. [ABSTRACT FROM AUTHOR]

Annual flower production ranged over four orders of magnitude among individuals of the tropical tree Prockia crucis (Flacourtiaceae), but the number of visits per flower by bees did not increase with flower number. In a population in Jalisco, México, the trees flower for about one week each year, offering only pollen to the bees (mostly solitary species). In a small group of trees, the number of visits per flower was less on trees with greater flower production but in a large group visitation did not vary between trees. Pollen flow probably was not directed from large to small trees or vice versa, because the number of flowers per tree did not determine the schedule of visits. The ratio of pollinators to pollen thieves decreased rapidly through the day, while individuals of both groups foraged more rapidly. Most bee species were rare, and only a small subset of medium-sized to large bees were effective pollinators. Large and small groups of trees differed in the relationship between individual flower crop and abundance and diversity of both pollinators and thieves. [ABSTRACT FROM AUTHOR]

(1) Background: Cardiocrinum cordatum (Thunb.) Makino (Liliaceae) is a forest perennial herb distributed in East Asia. Although flower visitors for this plant species have been well reported, their contribution to pollination remains unknown. (2) Methods: We evaluated pollination contribution for visitors of C. cordatum flowers in a natural cool temperate forest. We investigated visiting frequency, the number of pollen grains per body surface, fruit set, and the mean number of seeds per fruit produced after a single visit of each visiting species. Combining the results of these experiments, we determined the most important pollinators of this species. (3) Results: For the population investigated in the study, the three most essential pollinators were the bumblebee (Bombus diversus tersatus) (Apidae), sweat bee (Halictidae sp.), and marmalade hoverfly (Episyrphus balteatus) (Syrphidae). Additionally, we found that the contribution of a flower-visiting ant species (Myrmica ruginodis Nylander (s.l.)) (Formicidae) is small. (4) Conclusions: Pollinator contributions differed among flower visitors. Our results underscore the insufficiency of current information about flower-visiting species to evaluate pollination contribution. [ABSTRACT FROM AUTHOR]