The pollination biology ofHibbertia stricta (Dilleniaceae)

The exines of pollen grains ofHibbertia stricta (DC.)R. Br. exF. Muell. (Sect.Pleurandra) wear an oily, yellow pollen coat that stains positively for lipids. The pollen is collected by asocial bees, exclusively. The most common floral foragers are members of the genusLasioglossum (subgenusChilalictus;Halictidae) and they harvest pollen via thoracic vibration. As these bees cling to the inflated anthers their pollen smeared bodies come in contact with either of the two wet, nonpapillate stigmas. The stigmas respond positively to cytochemical tests for the presence of esterase immediately following expansion of the corolla, indicating the effective pollination period. The foraging patterns of the bees are narrowly to broadly polylectic. AsH. stricta flowers are nectarless, it is not surprising that bees bearing mixed pollen loads always carry the pollen of at least one nectariferous, coblooming plant. The pollination biology ofH. stricta is compared with otherHibbertia spp. and with pollen flowers in general.     

Bee-pollination inHibbertia fasciculata (Dilleniaceae)

In direct contrast to mostHibbertia spp., the flowers ofH. fasciculata R. Br. ex D. C. bear only a single whorl of stamens and these stamens are arranged separately (not in typical bundles). The short filaments are appressed to the three carpels so that the inflated, porose and introrsive anthers form a centralized cluster obscuring the three ovaries. The three slender styles emerge at right angles from between the filaments. These styles curve upward and the stigmas form the three points of a triangle; each stigma is approximately one millimeter outside the centralized cluster of anthers. The flowers are nectarless and bear a bright yellow corolla. A pungent and unpleasant fragrance appears to be concentrated within the pollenkitt. When native bees attempt to forage for the pollen, within the cluster of anthers, the ventrally deposited loads of pollen, on the bees' abdomens, contact the outer triangle of stigmas. The major pollinators ofH. fasciculata are female bees in the polylectic genera,Lasioglossum (subgenusChilalictus, Halictidae) andLeioproctus (Colletidae). These bees carry an average of more than two pollen taxa when they are caught foraging onH. fasciculata. 78% of the 47 bees, captured onH. fasciculata carried the pollen from at least one sympatric taxon bearing nectariferous flowers (e.g., genera in theMyrtaceae, Compositae, andEpacridaceae). The pollination biology ofH. fasciculata is assessed in relation to the known radiation of bee-pollinated flowers in the genusHibbertia, and within theDilleniaceae s. l.     

Pollination by a hugging mechanism inVigna vexillata (Leguminosae-Papilionoideae)

The pollination ofVigna vexillata (Leguminosae-Papilionoideae) by a carpenter bee,Xylocopa gualanensis (Hymenoptera-Anthophoridae) was studied in a secondary vegetation in Costa Rica. The bees were observed foraging onV. vexillata only in early mornings. Visits on individual flowers lasted about 7–8 seconds. Flower—pollen vector interactions are described and illustrated. By its pressure on the left-hand wing- and keel-petal in the asymmetrical flower, the weight of the bee causes the upper bearded part of the style along with the upper free parts of the stamens to slip out of the rigid keel-beak, hugging the bee over the dorsal part of its head and thorax. The occurrence of nototribic pollination inVigna and related genera is discussed.     

Bees assess pollen returns while sonicating Solanum flowers

Summary Can bees accurately gauge accumulating bodily pollen as they harvest pollen from flowers? Several recent reports conclude that bees fail to assess pollen harvest rates when foraging for nectar and pollen. A native nightshade (Solanum elaeagnifolium Cavanilles) that is visited exclusively for pollen by both solitary and social bees (eg. Ptiloglossa and Bombus) was studied in SE Arizona and SW New Mexico. The flowers have no nectaries. Two experiments were deployed that eliminated pollen feedback to the bees by experimentally manipulating flowers prior to bee visits. The two methods were 1) plugging poricidal anthers with glue and 2) emptying anthers of pollen by vibration prior to bee visitation. Both experiments demonstrated that bees directly assess pollen harvest on a flower-by-flower basis, and significantly tailor their handling times, number of vibratile buzzes per flower and grooming bouts according to the ongoing harvest on a given flower. In comparison to experimental flowers, floral handling times were extended for both Bombus and Ptiloglossa on virgin flowers. Greater numbers of intrafloral buzzes and numbers of times bees groomed pollen and packed it into their scopae while still on the flower were also more frequent at virgin versus experimental flowers. Flowers with glued andreocia received uniformly brief visits from Bombus and Ptiloglossa with fewer sonications and virtually no bouts of grooming. Curtailed handling with few buzzes and grooms also characterized visits to our manually harvested flowers wherein pollen was artificially depleted. Sonicating bees respond positively to pollen-feedback while harvesting from individual flowers, and therefore we expect them to adjust their harvesting tempo according to the currency of available pollen (standing crop) within Solanum floral patches.     

“Ixoroid” secondary pollen presentation and pollination by small moths in the Malagasy treeletIxora platythyrsa (Rubiaceae)

The pollination biology ofIxora platythyrsa (Rubiaceae) was studied in NW. Madagascar. The plant displayed cream-yellow, nocturnally fragrant, nectariferous, tubular and strongly protandrous flowers. These had an ixoroid secondary pollen presentation mechanism: prior to anthesis, anthers exhausted their pollen onto unripe stylar heads. From this position pollen of male-stage flowers later adhered to primarily the probosces of small visiting nocturnal noctuid and geometrid moths. — Pollen was subsequently raked off moths' probosces by receptive, copiously papillose stigmas of female-stage flowers. Principal pollination adaptation was probably to the noctuid moth subfam.Sarrothripinae.     

Bumble bee behavior and selection on flower size in the sky pilot,Polemonium viscosum

Summary In alpine Polemonium viscosum, plants having sweet-scented flowers are primarily pollinated by queens of the bumble bee species, Bombus kirbyellus. In this paper we ask whether two aspects of the pollination effectiveness of bumble bees, visitation rate and pollination efficiency, vary significantly with flower size in sweet-flowered P. viscosum.(i) Bumble bees visited plants with large flowers on 80–90% of encounters, but visited those with smaller flowers on only 49% of encounters. (ii) However, the gain in pollination that large-flowered plants obtained via increased visitation was countered in part because bumble bees deposited fewer outcross pollen grains per visit on stigmas of large flowers than on those of small ones. When both visitation rate and pollination efficiency are taken into account, the predicted value of a single bumble bee encounter declines from 1.06 seeds for flowers larger than 18 mm in diameter to 0.55 seeds for flowers smaller than 12 mm in diameter. Our results suggest that bumble bee pollinators of P. viscosum prefer flower morphologies that are poorly suited for precise pollination. Such behavioral complexities are likely to place constraints on the evolution of optimal floral design.     

Notes on the floral morphology in Vivianiaceae (Geraniales)

The functional floral morphology of the three genera of Vivianiaceae (= Ledocarpaceae, Geraniales), Rhynchotheca, Viviania and Balbisia, is compared. Likely pollination mechanisms are inferred from morphology and field observations. The flowers of Viviania are nectariferous and apparently zoophilous with nectar as the (primary) pollinator reward. Balbisia has pollen flowers without nectaries, its showy corolla indicates that it is also zoophilous with pollen as sole pollinator reward; bees were observed as flower visitors. One taxon (B. gracilis) may be anemophilous. Rhynchotheca has flowers without petals, with large, pendulous anthers and lacks nectaries. It shows synchronous mass flowering in its natural populations and is evidently anemophilous. A comparison with other Geraniales shows that nectar flowers with small anthers are likely the ancestral condition in Vivianiaceae. This suggests that the pollen flowers with larger anthers of Balbisia and Rhynchotheca may represent an apomorphic condition. The documentation of pollen flowers and anemophily in Vivianiaceae expands the range of known floral and pollination syndromes in Geraniales.     

Pollination system and its significance on isolation and hybridization in JapaneseEpimedium (Berberidaceae)

Observations on native populations of JapaneseEpimedium have revealed that two types of effective pollinators can be recognized. One of the two types, which consists of small bees (mainlyAndrena spp. andLasioglossum spp.), is characterized by nondiscriminating behavior for collecting pollen and is commonly found inEpimedium. The other type, which comprises medium sizedTetralonia nipponensis and largerBombus diversus queens as main components, showed flower-dependent foraging fidelity associated with nectar-sucking behavior.T. nipponensis with a shorter proboscis pollinated flowers with a shorter spur ofE. trifoliatobinatum and of a part ofE. s sempervirens, while the queen ofB. diversus with a longer proboscis pollinated longer spurred flowers ofE. grandiflorum andE. sempervirens. In the populations of putative hybrid-derivatives which show gradational variations of spur length, bees of the pollencollecting type pollinated any flower non-discriminately while bees of the nectar-foraging type tended to visit the flowers with spur lengths corresponding to their proboscis length. These observations suggest that the pollen-collecting bees play an important role for gene flow among theEpimedium species, and the nectar-foraging bees reinforce the isolation between the species by their selective pollination. Reproductive isolation between species ofEpimedium is discussed in relation with some practical behavior, such as flying power, of the pollinators.     

Pollination of Campanula rapunculus L. (Campanulaceae): How much pollen flows into pollination and into reproduction of oligolectic pollinators?

We studied an isolated population of Campanula rapunculus and two oligolectic bee species of Chelostoma (Megachilidae), their main pollinators. The population of C. rapunculus consisted of 2808 plants. Measurements of pollen flow showed that 3.7% of the pollen produced by a flower contribute to pollination, 95.5% was collected by bees for their offspring and 0.8% remained on the styles. Pollen analyses of brood cells of Chelostoma rapunculi revealed that females collected on average 4.9 million Campanula pollen to rear one bee. We calculated that approximately 1588 bees of this species could have been reared at the study site during the studied season. The amount of potentially viable pollen deposited on stigmas was 3.6 to 10.7 times higher than the number of ovules. We discuss morphological features of the flowers which may lower the pollen removal rate per bee visit and consequently cause a high visitation and pollination rate.The study was supported by a joint project CAPES / DAAD (Probral 112/00). We thank NaBu (Naturschutzbund) Bonn and the Untere Landschaftsbehörde Siegburg for the permission to work at the Natural Reserve Dünstekoven.     

Competition between the oligolectic beePtilothrix plumata (Anthophoridae) and the flower closing beetlePristimerus calcaratus (Curculionidae) for floral resources ofPavonia cancellata (Malvaceae)

The flowers ofPavonia cancellata, a creeping ruderal half-shrub of northeastern Brazil, open synchronously at 6:00 h with all anthers already dehisced. The oligolectic beePtilothrix plumata was the most effective pollinator. During 90—180 min, female bees make up to 40 brief pollen collection trips to provision their brood cells. The pollen of about 40 flowers ofP. cancellata is needed to feed one bee larva. The most frequent flower visitors, however, are the specialized curculionid beetlesPristimerus calcaratus, which do not crosspollinate the flowers. They perforate the epidermis with their mouthparts, provoking dehydration, and then actively close the loose petals with their legs. Two hours after opening, half of the flowers had already been closed by the beetles. We interpret the fast, uninterrupted pollen foraging ofPtilothrix plumata bees as a strategy adapted to synchronous pollen presentation ofPavonia and to competition withPristimerus calcaratus: the female bees have to provision their brood cells before the beetles succeed in closing the flowers.     

Floral mimesis inThelymitra nuda (Orchidaceae)

Insect pollination occurred inThelymitra nuda R. Br. on sunny days when the ambient temperature exceeded 20 °C. The flower buds on a raceme opened subsynchronously displaying the brightly-colored, actinomorphic perianth and exposed the contrastingly-colored, scented and ornamented column. InT. nuda the staminodes and the filament of the fertile stamen are fused to one another producing an inflated hood over the anther. This staminodal hood is terminated in two non-ornamental, but brightly-colored, central lobes and two terete lateral lobes bearing approximately 400 white trichomes. Each trichome bears a double chain of 30–40 spherical, rugulose cells. Female bees, in the genusLasioglossum (Halictidae), were observed to land directly on the hood and curled their bodies around the four lobed tip of the staminodal complex. The bees attempted to forage on the lobes as if they were collecting pollen from fertile, poricidal stamens. These bees applied thoracic vibration to the yellowish central lobes and actively scraped the trichome clusters (Pseudopollen) with their forelegs. Bees carried the pollinaria ofT. nuda dorsally on their abdomens. Abdominal contact with the rostellum appeared to occur when the female bee disengaged herself from the staminodal hood. Observations made of bees on co-blooming flowers, and analyses of pollen loads collected by bees suggested that the orchid flower mimics the guild of blue-purple flowers that lack floral nectar but offer pollen in poricidal anthers. The models ofT. nuda are co-bloomingLiliaceae in the generaDichopogon andThysanotus. However, nectarless, buzz-pollinated flowers are also extensively distributed over the orchid’s range. More than 30% of the flowers in theT. nuda population had their pollinaria removed, suggesting a high capacity for cross-pollination in an orchid genus usually considered to be self-pollinated via mechanical autogamy. This study confirmed previous predictions that column modifications represented a trend towards pseudanthery.     

Pollination biology ofSymphonia globulifera (Clusiaceae)

The pollination biology ofSymphonia globulifera was studied in Central Amazonia, Brazil. As suggested by the bird syndrome of the flowers, these are mainly pollinated by hummingbirds. Occasional visits by other birds, butterflies and more rarely bees, as well as tamarin monkeys were also observed.Trigona bees partly destroy the flower tube to rob nectar. The possibility thatS. globulifera may not be primarily adapted to hummingbird pollination is discussed. The pollen is intermixed in an oily fluid secreted by the anthers (antheroil). Each of the five stigmas consists of a pore-like opening at the apex and a small chamber behind it. The antheroil mixed with pollen is absorbed by capillarity into the chamber when deposited on the pore. the pollen germinates inside the stigma. The presence of antheroil and pore-like stigmas in the flowers of the closely relatedPlatonia insignis indicate a similar mode of pollination. The results of this study are compared with observations in some otherClusiaceae (Caraipa, Clusia, Garcinia, Mahurea), where floral oils or floral resin occur. The role of these substances in the pollination process and their relation to the evolution of flower biology inClusiaceae are briefly discussed.     

Anthecology ofSchrankia nuttallii (Mimosaceae) on the tallgrass prairie

Schrankia nuttalii flowers through late spring on the tallgrass prairie. Although each stem produces an average of 26 capitate inflorescences only 12% of those inflorescences will open each day to disperse and receive polyads. Each inflorescence may live up to 48 hours but anthers abscise by late afternoon on the first day and the filaments change color and lose their scent. The 78–93 florets comprising each inflorescence open synchronously before dawn or during early morning hours. First day inflorescences ofS. nuttallii are herkogamous and fragrant. They are nectarless. Bombyliid flies and male bees are infrequent floral foragers so the major pollinators include female bees representing five families;Anthophoridae, Apidae, Colletidae, Halictidae, andMegachilidae. All foraging insects ignore second day inflorescences although stigmas are still receptive. Although 97% of all bees collected onS. nuttallii carrySchrankia polyads in their scopae or corbiculae 59% also carry the pollen/pollinaria of one or more coblooming angiosperms. At least 98% of all bees carrying mixed pollen loads incorporate the pollen/pollinaria of one or more nectariferous taxa (e.g.Asclepias spp.,Asteraceae, Convolvulaceae, Delphinium spec., etc.). Species of halictid bees are more likely to carry pure loads ofS. nuttallii polyads (70%) than bees of the four remaining families. Due to the nectarless florets and high degree of polylectic foraging bee-pollination inS. nuttallii converges more closely with the pollination systems of some AustralianAcacia spp. than with most other xeric/tropical genera of mimosoids studied in the western hemisphere.     

Different responses of pollinating bees to size variation and sexual phases in flowers ofCampanula

To examine the response of pollinating bees to size and sexual phases of flowers, we constructed an artificial population ofCampanula having large flower variation and presented it to potentially pollinating bees in nurseries. The pollinating bee groups (halictid, megachilid and bumble bees) responded differentially to both the flower size and to the sexual phases of the flowers. Whereas visitation rate of megachilid bees increased with the flower size, those of halictid bees and bumble bees did not show particular trends; for example, bumble bees visited almost all of the flowers consistently. Visitation frequencies to male-and female-phased flowers were significantly different between megachilids at Tokyo and halictids. This study indicates that pollinator attraction could not solely explain the evolution of the flower size inCampanula, and that other factors such as pollen transfer efficiency, should be considered.     

Stamen movements in flowers ofOpuntia (Cactaceae) favour oligolectic pollinators

Opuntia brunneogemmia andO. viridirubra occur sympatrically in the Serra do Sudeste, Rio Grande do Sul, Brazil. Their flowers have 450–600 thigmonastic stamens and provide large amounts of pollen and nectar for bees. Bees of 41 species were registered at the flowers ofO. brunneogemmia and 30 at the flowers ofO. viridirubra. Females of three oligolectic species are the only effective pollinators:Ptilothrix fructifera (Anthophoridae),Lithurgus rufiventris (Megachilidae), andCephalocolletes rugata (Colletidae). During their visits inOpuntia-flowers, bees touch the filaments and stimulate the movement of the stamens to the centre of the flower. At the end of this movement, the anthers are densely packed around the style. As a consequence the pollen is presented in an easily accessible upper layer of anthers and various, nearly inaccessible lower layers. The lower layers contain about 80% of the pollen reward. Only females of the three oligolectic pollinators exploit the pollen from the lower layers and reach the nectar furrow. Therefore, through their stamen movements,Opuntia flowers hide most of their pollen from flower visitors but favour effectively pollinating, oligolectic bees.     

The floral biology ofThelymitra epipactoides (Orchidaceae), and the implications of pollination by deceit on the survival of this rare orchid

Thelymitra epipactoides has a highly variable visual display achieved through polychromatic flowers and variable inflorescence size, bearing between 7 and 31 flowers, which attract foraging polylectic bees. Only bees of the genusNomia were observed carrying pollinia and successfully pollinating the orchid. The genusNomia contains polylectic, pollen gathering species that store pollen in both the crop and scopa on the hind legs. The absence of a reward for the bees indicates the orchid is relying on deception to attract visitors. The relationship of deception to mimicry is discussed. Once on the flower, tactile, visual and possibly olfactory stimuli direct bees to the false anther formed by the voluminous column wings, where morphological adaptations of the flower ensure that the pollinarium is deposited on the gaster of the bee to effect pollination. — The lack of seed set observed on the Victorian coast appears to be due to the absence of pollinators from the heath and grassland communities in which the orchid grows. This may well be a consequence of the reduced number of plants flowering in the community (a result of the elimination of fire at these sites), thus not maintaining a floral community attractive to potential pollinators.     

Effectiveness of honey bees in delivering the biocontrol agent Bacillus subtilis to blueberry flowers to suppress mummy berry disease

Honey bees are important pollinators of commercial blueberries in the southeastern United States, and blueberry producers often use supplemental bees to achieve adequate fruit set. However, honey bees also vector the plant pathogenic fungus Monilinia vaccinii-corymbosi which infects open blueberry flowers through the gynoecial pathway causing mummy berry disease. Here, we report the results of a 3-year field study to test the hypothesis that using bee hives equipped with dispensers containing the biocontrol product Serenade, a commercial formulation of the bacterium Bacillus subtilis which has shown activity against flower infection by M. vaccinii-corymbosi in laboratory experiments, can reduce mummy berry disease incidence when honey bees are used as pollinators in blueberries. Individual honey bees carried 5.1–6.4 × 10⁵ colony-forming units (CFU) of B. subtilis when exiting hive-mounted dispensers with Serenade. On caged rabbiteye blueberry bushes in the field, population densities of B. subtilis vectored by honey bees reached a carrying capacity of <10³ CFU per flower stigma within 2 days of exposure, and there was a highly significant non-linear relationship between B. subtilis populations per stigma and bee activity, expressed as number of legitimate flower visits per time interval per cage (R = 0.6928, P < 0.0001, n = 32). Honey bee density (1600 or 6400 individuals per 5.8-m³ cage) and Serenade treatment (presence or absence of the product in hive-mounted dispensers) significantly (P < 0.05) affected the incidence of fruit mummification on caged bushes, whereby increasing bee density increased disease incidence and application of Serenade reduced disease levels. Taken together, results of this study suggest that use of a hive-dispersed biocontrol product such as Serenade as a supplement during pollination can reduce the risk of mummy berry disease. This may be a prudent practice that optimizes the benefits to pollination of high bee densities while reducing the associated disease-vectoring risk.     

The effectiveness of heterostyly in preventing illegitimate pollination in dish-shaped flowers

Heterostyly is thought to have evolved because it confers efficiency of cross-pollination. While it can be an alternative to an incompatibility mechanism in tube-shaped flowers, its effectiveness in dish-shaped flowers, such as those of Fagopyrum esculentum, has not been demonstrated. Stigmatic pollen loads were examined over the daily course of pollination in a planted field of this crop species. Natural pollination by bees resulted in 23% of the pollen on pin flowers and 69% on thrum flowers being legitimate. Both flower morphs therefore received the two pollen types in about the same proportions. Pin pollen constituted 74% of the pollen delivered to stigmata, even though the two morphs are present in equal numbers. Pin flowers collected about 75 pollen, and thrum flowers about 25 pollen by the end of the day of anthesis. The net result was about 15 compatible pollen per flower on both floral morphs. These results indicate that heterostyly is associated with excess pin-pollen delivery rather than excess legitimate-pollen delivery in F. esculentum.     

The internal transcribed spacer 2 region of the nuclear ribosomal DNA and the phylogeny of the moss family Hylocomiaceae

Eichhornia azurea is a tristylous species of Pontederiaceae. Moderate self-incompatibility accompanies the floral heteromorphism of this species.Eichhornia azurea is almost exclusively visited by one bee species,Ancyloscelis gigas (Anthophoridae), in South Brazil. This species has an extremely long proboscis covered with recurved hairs. Bees use these hairs to collect pollen from the anthers placed inside the narrow perianth tube ofE. azurea. Analyses of the pollen load ofA. gigas females show that they are flower constant and carry pollen from all three anther levels. The behavior ofA. gigas and its proboscis morphology make this bee species narrowly adapted and an effective pollinator for facilitating legitimate pollination ofE. azurea flowers in southern Brazil.     

Pollination of four sympatric species ofAngelonia (Scrophulariaceae) by oil-collecting bees in NE. Brazil

The manner whereby the oil-producing bisaccate flowers ofAngelonia (Scrophulariaceae) are pollinated by female oil-collecting bees is reported for the first time. Observations were made in the Caatinga formation of Pernambuco, NE. Brazil, on four synchronopatric species. These differ in sizes and structural details of the corolla, level of flower exposition, and habitat preferences. All legitimate visitors wereCentris spp. (Anthophoridae):Angelonia hirta was mainly pollinated byC. fuscata andA. pubescens byC. hyptidis; A. bisaccata andA. hookeriana shared an unidentified species. Several exomalopsine, tetrapediine and meliponid bees exploit the flowers less descriminately for oil or pollen, respectively, without regularly contacting anthers and stigma. The flowers are protandrous, and are self-incompatible except those of the annualA. pubescens. After alighting, theCentris bees introduce their front legs simultaneously into each of the pouches and start alternate collecting movements to gather the oil from the trichome elaiophores. While doing so, they are forced by projections of the corolla floor to press their head under the anthers and stigma, whereby pollen is transferred with their frons or clypeus. On account of their collector type and behaviour,C. fuscata andC. spec. are not specialized toAngelonia but may equally exploit other nonrelated taxa for oil, whereasC. hyptidis exhibits oligolecty onA. pubescens. It possesses relatively elongate forelegs with padlike collectors suitable for sweeping the lipids from the scattered glandular hairs inside the divergent spurs of its host. It is the only species that also collects pollen (by buzzing) from the oil host.A. hirta and relatives, provided with dense elaiophore carpets, are, for their part, adapted to scrapingCentris species with typical oil collectors. Flower and bee phenologies, although largely dependent on the irregular rainfalls, are not always coincident.