The effects of landscape on bumblebees to ensure crop pollination in the highland agricultural ecosystems in China

Honey bees and wild bees provide critical pollination services to agricultural ecosystems; however, the relative contributions of different bee taxa are not well understood. The natural habitats surrounding farmland support food and nesting resources for wild bees and therefore play an important role in the maintenance of crop pollination. In this study, we selected Cucurbita pepo L. (squash) as a model crop to investigate the relative importance of honey bees and bumblebees in pollinating the crop. Thirteen fields, which were surrounded by a gradient of natural habitat, were investigated on the Yunnan‐Guizhou Plateau in China. We measured the visit densities of honey bees and bumblebees, the number of pollen grains deposited in a single visit by the two bee taxa, as well as the overall pollen grains deposited on stigmas during a flowering day, and then used Bayesian inference to decouple the pollen grains deposited by either the honey bees or the bumblebees. Compared with honey bees, bumblebees deposited a higher number of pollen grains on stigmas in a single visit, but had a lower visit density than honey bees. Meanwhile, the bumblebee visit density increased along the proportion of natural habitat, while the honey bee visit density was not affected by the surrounding natural habitat. Data simulations using Bayesian inference showed that on a flowering day, the number of pollen grains deposited by bumblebees increased with the proportion of natural habitat in the surrounding landscape, but the number of pollen grains deposited by honey bees did not. Moreover, the total numbers of pollen grains deposited by honey bees or bumblebees alone were all below 2000 (the critical level to satisfy the pollination requirement of this crop). Pollen calculations demonstrated that the number of pollen grains deposited by the two bee taxa was greater than 2000 in fields surrounded by more than 13% natural habitat (grasslands and forests). The results revealed that bumblebees ensured C. pepo pollination in combination with honey bees in the highland agricultural ecosystems.     

Foraging behavior affects pollen removal and deposition in Impatiens capensis (Balsaminaceae)

Flowers of most plant species are visited by a variety of animals. Some of these visitors are effective pollinators while others remove resources without transferring pollen. Studies comparing the effectiveness of different visitors as pollinators often compare taxa without considering variation in behavior within a taxon. Wilson and Thomson (Ecology 72: 1503-1507, 1991) documented the effects of honey bees and bumble bees on the pollination dynamics of Impatiens capensis. They found that pollen-collecting honey bees removed large numbers of pollen grains from anthers but deposited little of it on stigmas; bumble bees, which sought nectar, removed less pollen but deposited more of it on stigmas. It is unclear whether the low pollen transfer efficiencies of honey bees are explained by their morphology or by their pollen-collecting behavior. We repeated the work of Wilson and Thomson at a site where honey bees were foraging for nectar, not pollen. We measured the quantity of pollen remaining in anthers, the number of pollen grains deposited on stigmas, and seed production after single visits by honey bees and bumble bees. The differences between the taxa disappeared when they were foraging in a similar manner. Our results clearly demonstrate the importance of foraging behavior on the pollination effectiveness of floral visitors.     

Floral resources foraged by Geotrigona argentina (Apidae,Meliponini) in the Argentine Dry Chaco forest

This study is the first contribution to knowledge of the relationships between Geotrigona argentina and the plants of the Argentine Dry Chaco forest. A total of 1260 g of honey (corresponding to 146 pots) and 763 g of pollen (63 pots) stored in four underground nests was studied. The honey pots from each nest were homogenised and the four honey samples were analysed by melissopalynological methods, whereas the pollen pots were studied individually. Both classical counts and counts affected by the volume of the pollen types were carried out. Pollen data were statistically analysed. Additional data on both protein and lipid content is also provided. A total of 39 pollen taxa were identified. Pollen collection was focused on a few pollen taxa: Prosopis, Castela coccinea, Maytenus and Capparis; these taxa, together with Ziziphus mistol and Pisonia zapallo, were also important nectar sources. The preliminary results show that pollen collection varied seasonally, being most diverse in the summer when G. argentina incorporates herbaceous plants into its diet. The pollen collection spectrum of G. argentina is similar to that of other Trigonina bees in that the main plant species collected are a few large shrubs or trees, whose flowering consists of small and clustered flowers. Pots with large amounts of monofloral loads with pollen from only a few species suggests an organised foraging behaviour that includes the recruitment of foragers, such as that observed in other eusocial bees.     

The effects of natural variation in pollinator visitation on rates of pollen removal in wild radish,Raphanus raphanistrum (Brassicaceae)

Pollen removal from flowers is an important component of male fitness, but the effects of natural variation in visitation rates on pollen removal are poorly understood. We measured pollen removal over 2 yr in experimental field populations of wild radish, Raphanus raphanistrum. Pollen removal and pollinator visitation over 1-hr periods were measured on previously unvisited flowers. The effects of pollen production and visitation by different insect taxa on pollen removal were determined using multiple regression. Pollen removal rates were extremely high; a median of 84% of pollen produced was removed in 1 hr. Pollen production was far more important than visitation in determining the number of pollen grains removed. Pollen removal increased with increasing numbers of visits by honey bees and small native bees, but increased numbers of syrphid fly visits had no effect. Average visit duration had no effect on pollen removal in 1991, and a marginally negative effect in 1992.     

Pollinator genetics and pollination: do honey bee colonies selected for pollen‐hoarding field better pollinators of cranberry Vaccinium macrocarpon?

1. Genetic polymorphisms of flowering plants can influence pollinator foraging but it is not known whether heritable foraging polymorphisms of pollinators influence their pollination efficacies. Honey bees Apis mellifera L. visit cranberry flowers for nectar but rarely for pollen when alternative preferred flowers grow nearby. 2. Cranberry flowers visited once by pollen‐foraging honey bees received four‐fold more stigmatic pollen than flowers visited by mere nectar‐foragers (excluding nectar thieves). Manual greenhouse pollinations with fixed numbers of pollen tetrads (0, 2, 4, 8, 16, 32) achieved maximal fruit set with just eight pollen tetrads. Pollen‐foraging honey bees yielded a calculated 63% more berries than equal numbers of non‐thieving nectar‐foragers, even though both classes of forager made stigmatic contact. 3. Colonies headed by queens of a pollen‐hoarding genotype fielded significantly more pollen‐foraging trips than standard commercial genotypes, as did hives fitted with permanently engaged pollen traps or colonies containing more larvae. Pollen‐hoarding colonies together brought back twice as many cranberry pollen loads as control colonies, which was marginally significant despite marked daily variation in the proportion of collected pollen that was cranberry. 4. Caloric supplementation of matched, paired colonies failed to enhance pollen foraging despite the meagre nectar yields of individual cranberry flowers. 5. Heritable behavioural polymorphisms of the honey bee, such as pollen‐hoarding, can enhance fruit and seed set by a floral host (e.g. cranberry), but only if more preferred pollen hosts are absent or rare. Otherwise, honey bees' broad polylecty, flight range, and daily idiosyncrasies in floral fidelity will obscure specific pollen‐foraging differences at a given floral host, even among paired colonies in a seemingly uniform agricultural setting.     

Nectar and Pollen Sources for Honeybee （Apis cerana cerana Fabr.） in Qinglan Mangrove Area, Hainan Island, China

In the present study, nectar and pollen sources for honeybee （Apls cerana cerana Fabr.） were studied in Qlnglan mangrove area, Hainan Island, China, based on microscopic analysis of honey and pollen load （corblcular and gut contents） from honeybees collected In October and November 2004. Qualitative and quantitative melittopalynologlcal analysis of the natural honey sample showed that the honey is of unlfloral type with Mimosa pudlca L. （Mlmosaceae） as the predominant （89.14%） source of nectar and pollen for A. cerana cerana In October. Members of Araceae are an Important minor （3%-15%） pollen type, whereas those of Arecaceae are a minor （〈3%） pollen type. Pollen grains of Nypa fruticans Wurmb., Rhlzophora spp., Excoecarla agallocha L., Lumnitzera spp., Brugulera spp., Kandella candel Druce, and Ceriops tagal （Perr.） C. B. Rob. are among the notable mangrove texa growing In Qinglan mangrove area recorded as minor taxa In the honey. The absolute pollen count （I.e. the number of pollen grains/10 g honey sample） suggests that the honey belongs to Group V （〉1 000 000）. Pollen analysis from the corblcular and gut contents of A. cerana cerana revealed the highest representation （95.60%） of members of Sonneratia spp. （Sonneratlaceae）, followed by Bruguiera spp. （Rhizophoraceae）, Euphorblaceae, Poaceae, Fabaceae, Arecaceae, Araceae, Anacardlaceae, and Rublaceae. Of these plants, those belonging to Sonneratla plants are the most Important nectar and pollen sources for A. cerana cerana and are frequently foraged and pollinated by these bees in November.     

Seasonal patterns of foraging activity in colonies of the African honey bee,Apis mellifera scutellata,in Africa

Summary Seasonal foraging patterns were investigated using six observation colonies maintained in the Okavango Delta, Botswana. Pollen collection, flight from the hive, and recruitment for pollen and nectar sources occurred throughout the 11 months of the study. However, the distribution of foraging activity throughout the day changed seasonally. Colonies emphasized recruitment for pollen sites throughout most of the year. Brood production occurred in all months except May, and there was a significant, positive correlation between the proportion of recruitment activity devoted to pollen sources and the amount of brood comb in the colonies. The seasonal foraging patterns ofscutellata in the Okavango were similar to those of Africanized honey bees in the neotropics. The extended foraging season and emphasis on pollen collection may be associated with the high swarming rates and migrational movements of tropical honey bees.     

The effect of wild radish floral morphology on pollination efficiency by four taxa of pollinators

The effects of floral morphology on rates of pollen removal and deposition by different pollinators in generalist plant species are not well known. We studied pollination dynamics in wild radish, Raphanus raphanistrum, a plant visited by four groups of pollinators: honey bees, small native bees, butterflies, and syrphyd flies. The effects of anther position and other factors on pollen removal during single visits by all four pollinator taxa were measured. Flowers with high anther exsertion (i.e., anthers placed higher above the opening of the corolla tube) tended to have the highest numbers of pollen grains removed, but this effect was strongest for honey bees and butterflies. For all pollinator taxa, pollen removal increased with the number of pollen grains available on a flower and whowed a positive, decelerating relationship with the duration of the visit. The effects of stigma position and other factors on pollen deposition during single visits by honey bees and butterflies were also studied. The nectar-feeding butterflies had a higher pollination efficiency (percentage of pollen grains removed from anthers that were subsequently deposited on a stigma) than the nectar- and pollen-feeding honey bees. Flowers with intermediate stigma exsertion had the highest numbers of pollen grains deposited on their stigmas by butterflies, but stigma exsertion had no effect on deposition by honey bees. For both butterflies and honey bees, pollen deposition on the recipient flower increased with the amount of pollen removed from the donor flower, and there was a positive, decelerating relationship between deposition and time spent at the flower; these results are analogous to those for pollen removal. The effects of anther and stigma exsertion on pollen removal and denosition did not fit predictions based on patterns of floral correlations, but results for morphology, pollen availability, time spent per visit, and pollinator efficiency are in broad agreement with previous studies, suggesting the possible emergence of some general rules of pollen transfer.     

Pollen and yeast change nectar aroma and nutritional content alone and together,but honey bee foraging reflects only the avoidance of yeast

Floral nectar often contains pollen and microorganisms, which may change nectar's chemical composition, and in turn impact pollinator affinity. However, their individual and combined effects remain understudied. Here, we examined the impacts of the nectar specialist yeast, Metschnikowia reukaufii, and the addition of sunflower (Hellianthus annus) pollen. Pollen grains remained intact, yet still increased yeast growth and amino acid concentrations in nectar, whereas yeast depleted amino acids. Pollen, but not yeast, changed nectar sugar concentrations by converting sucrose to its monomers. Both pollen and yeast contributed emissions from nectar, though yeast volatiles were more abundant than pollen volatiles. Yeast volatile emission was positively correlated with pollen concentration and cell density, and yeast depleted a subset of pollen-derived volatiles. Honey bees avoided foraging on yeast-inoculated nectar and foraged equally among uninoculated nectars regardless of pollen content, underscoring the importance of microbial metabolites in mediating pollinator foraging.     

Seasonal variation in vegetation and pollen collected by honeybees in Thessaloniki,Greece

Pollen is important for the nutrition of honeybees and it is necessary for their survival and reproduction. In this study, we collected daily the pollen pellets from four colonies and also recorded the plants in flower in the area around the apiary, over a two‐year period. Field records revealed the presence of 204 species with Asteraceae, Fabaceae and Rosaceae being the most specious families. Although honey bees collected more than 140 pollen types, the main pollen sources (>60% of the total weight) came from less than ten taxa. The most important pollen types with respect to total weight were Sisymbrium irio, Papaver rhoeas, Verbascum sp., Polygonum aviculare, Zea mays and Olea europaea. The use of pollen traps proved a more accurate method to record the type and the foraging period during which the honeybees collect pollen, compared to field observations.     

Pollination biology in Roepera (Zygophyllaceae): How flower structure and shape influence foraging activity

The foraging behavior of bees is a complex phenomenon that depends on numerous physical features of flowers. Of particular importance are accessibility of floral rewards, floral proportions, symmetry and orientation. The flowers of Roepera are characterized by the presence of staminal scales (SS), which play an important role in nectar protection. We studied two species of Roepera with different symmetry and flower orientation, which are mainly visited by honeybees (Apis mellifera). We aimed to show how the foraging behavior of honey bees is affected by the function of SS, floral symmetry and orientation. The foraging behavior was documented by video photography. Handling time, access to nectar, percentage of pollen/nectar foraging, percentage of pollen contact and pollen deposition site on the honey bee's body were assessed. The morphometric features of the honey bees and flowers were analyzed. We found that the SS restricted pollinator access to nectar. Our results indicated consistency of visitation patterns in zygomorphic, laterally oriented flowers of R. fuscata versus random patterns in actinomorphic, diversely oriented flowers of R. leptopetala. The relative proportions of SS and proboscis length appear to be crucial for the success of pollinators. The directionality of the honey bees' movement, together with the different positioning of reproductive organs, plays an important role in the accuracy of pollen transfer and pollination efficiency.     

Seasonal variation of collected pollen loads of honeybees (Apis mellifera L. anatoliaca)

Pollen collected by honeybees foraging in the region of Bursa, Turkey was analysed for a whole year. Pollen loads were collected from the hives of Apis mellifera anatoliaca once a week and were classified by colour. Forty‐one taxa were identified from the pollen analyses of the loads and 14 of these had percentages higher than 1%. Only 2.05% of the total pollen could not have been identified. Dominant taxa include; Brassicaceae (11.19%), Helianthus annuus L. (10.84%), Cichorioideae (8.93%) Salix spp. (7.99%), Rosaceae (7.37%), Centaurea spp. (7.56%), Papaver spp. (7.41%), Knautia spp. (6.99%), Fabaceae (6.01%), Asteraceae (5.73%), Xanthium spp. (2.65%), Chrozophora spp. (2.45%), Plantago spp. (1.56%) and Acer spp. (1.54%) representing 88.23% of the total. Distinct variations in plant usage are seen through the year with initial use of Rosaceae, Salix, and to a lesser extent Brassicaeae. As these groups finish flowering the bees move onto Helianthus annuus, Centaurea through the summer followed by Asteraceae in the late summer and Fabaceae in the autumn. There is a strong reliance on crop species for pollen forage but a number of indigenous species are also seen within the samples. The most productive period for collecting various pollen types, and the ideal period to determine pollen preferences of honey bees was June‐August.     

Could learning of pollen odours by honey bees (Apis mellifera) play a role in their foraging behaviour?

Abstract. The role of pollen odour cues in the foraging behaviour of honey bees (Apis mellifera L.) is poorly understood. Using classical conditioning of the proboscis extension response, in which bees learn to associate an odour with a sucrose reward, the present study tests whether odours of bee-collected pollen from the hive environment or odours of fresh pollen on the anthers of flowers could be used in pollen foraging. Honey bees efficiently learn odours from field-bean (Vicia faba) bee-collected pollen and oilseed-rape (Brassica napus) bee-collected pollen, hand-collected pollen, anthers and whole flowers, demonstrating that honey bees can learn pollen odours associatively in biologically realistic concentrations. Honey bees learn pollen odours of oilseed rape better than field bean and, although they generalize these two odours, they easily distinguish between them in discrimination tests, suggesting that pollen odours may be used in species recognition/discrimination. There is little evidence that honey bees can recognize whole flowers based on previous experience of bee-collected pollen odour. However, they generalize the odours of oilseed-rape anthers and whole flowers, suggesting that anther pollen in situ may play a more prominent role than bee-collected pollen in foraging behaviour.     

Honey bee‐collected pollen in agro‐ecosystems reveals diet diversity,diet quality,and pesticide exposure

European honey bees Apis mellifera are important commercial pollinators that have suffered greater than normal overwintering losses since 2007 in North America and Europe. Contributing factors likely include a combination of parasites, pesticides, and poor nutrition. We examined diet diversity, diet nutritional quality, and pesticides in honey bee‐collected pollen from commercial colonies in the Canadian Maritime Provinces in spring and summer 2011. We sampled pollen collected by honey bees at colonies in four site types: apple orchards, blueberry fields, cranberry bogs, and fallow fields. Proportion of honey bee‐collected pollen from crop versus noncrop flowers was high in apple, very low in blueberry, and low in cranberry sites. Pollen nutritional value tended to be relatively good from apple and cranberry sites and poor from blueberry and fallow sites. Floral surveys ranked, from highest to lowest in diversity, fallow, cranberry, apple, and blueberry sites. Pesticide diversity in honey bee‐collected pollen was high from apple and blueberry sites and low from cranberry and fallow sites. Four different neonicotinoid pesticides were detected, but neither these nor any other pesticides were at or above LD₅₀ levels. Pollen hazard quotients were highest in apple and blueberry sites and lowest in fallow sites. Pollen hazard quotients were also negatively correlated with the number of flower taxa detected in surveys. Results reveal differences among site types in diet diversity, diet quality, and pesticide exposure that are informative for improving honey bee and land agro‐ecosystem management.     

Pollen of Apis honey from Maharashtra

Melissopalynological investigations are important in establishing an apiary industry. Pollen analyses of the nectar sources of Apis florea Fabr. from the Saoner region indicate the importance of this region for commercial bee keeping enterprises. Twenty seven samples were collected during winter season in the months of October 1997 to January 1998 and November 1998 to January 1999. In addition, 245 pollen loads were analyzed from 19 honey combs collected from the same region. Pollen grains were identified from reference slides. Of the 27 honey samples, 17 samples were unifloral and ten multifloral. A total of 39 pollen types were classified into 21 families. In the analyses of the 245 pollen loads Lagascea mollis Cav. was dominant in 57 pollen loads and Parthenium hysterophorus L. in 55 pollen loads. Weeds such as Parthenium hysterophorus, Sphaeranthus indicus L., Lagascea mollis, Bidens pilosa L. and Justicia procumbens L. were the common sources of nectar and pollen for Apis florea in the region. Agricultural tracts of the Saoner region in the Nagpur District have a good potential for sustaining beekeeping ventures because of the diversity of nectar and pollen taxa.     

Economic motivation for plant species preferences of pollen-collecting bumble bees

1. Pollen is an essential resource for bees as it provides the proteins needed for larval development and growth. Bumble bees typically collect pollen from a limited subset of the available plant species, indicating that bees perceive qualitative and/or quantitative differences between species. Because bumble bees typically exhibit different foraging behaviours when collecting pollen than when collecting nectar, and often visit different plant species, the economics of pollen collection are subject to unique benefits and costs.
2. The influences of pollen standing crop, grain volume, protein content and foraging costs on the choice of plant species by pollen-collecting bumble bees were assessed. Pollen was collected from different plant species available to bees and these species were ranked according to eleven currencies that differentially incorporate these influences on foraging economics.
3. Comparison of observed preferences at three sites with species rankings based on foraging currencies revealed that bumble bees do not assess plant species based solely on intrafloral characteristics. Instead, pollen-foraging bumble bees are sensitive to protein availability at the site as a whole as well as foraging costs, and forage in a manner that maximizes the site-specific efficiency of protein collection (protein collected/energetic costs). Such behaviour would promote proper larval development while maximizing forager lifespan.     

POLLINATION SYSTEMS IN PASPALUM DILATATUM POIR. (POACEAE): AN EXAMPLE OF INSECT POLLINATION IN A TEMPERATE GRASS

The pollination of Paspalum dilatatum was studied in south-central Oklahoma during the summer of 1979. Pollen was liberated between 0700–0900 hr except on humid mornings (RH ≤ 80%), when there was a delay of 2–3 hr and a reduction in the total air-borne pollen concentration. A rapid decrease in air-borne pollen concentration with distance from the source results from: 1) individual pollen grains larger (50–70 μ in diam) than typical wind-pollinated plants, and 2) some pollen dispersed as clusters of grains. Several floral characters of P. dilatatum led to a hypothesis that this perennial grass may be entomophilous as well as anemophilous. Not only were the pollen grains larger than most other wind-pollinated taxa but the species produced fewer than 2,500 pollen grains per ovule. Three additional observations corroborate this view: 1) solitary bees (Halictidae) actively collect pollen during the morning, 2) the number of pollen grains per stigma was significantly (P < 0.001) greater on racemes exposed to both wind and bees than on racemes exposed only to wind, and 3) the combination of bees and wind as pollinators significantly (P < 0.001) increased seed set compared to wind alone.     

Pollen analyses of honey from Finland

The present study reports results of qualitative melissopalynological analyses of Finnish honey between the years 2000–2007 and changes in its pollen content from the period 1960–2007. Altogether the pollen content of 734 honey samples was analysed with an average of 415 pollen grains counted from a sample. Pollen of Trifolium repens type, Rubus spp., Salix spp. and the Brassicaceae family were present in more than 90% of the samples, and these pollen types were also found in the highest proportions. Annual variation in the relative amounts of the most numerous pollen types could be as high as 10%. On the basis of the pollen spectra of the honey samples, four regions of forage plants for bees could be identified in Finland. In the period between 1960 and 2007, the most marked change observed was that the percentage of the Trifolium spp. pollen type had decreased from 70% to 10%, while the proportions of Brassicaceae and Rosaceae pollen types showed a corresponding increase.     

Honey bees and wild pollinators differ in their preference for and use of introduced floral resources

Introduced plants may be important foraging resources for honey bees and wild pollinators, but how often and why pollinators visit introduced plants across an entire plant community is not well understood. Understanding the importance of introduced plants for pollinators could help guide management of these plants and conservation of pollinator habitat. We assessed how floral abundance and pollinator preference influence pollinator visitation rate and diversity on 30 introduced versus 24 native plants in central New York. Honey bees visited introduced and native plants at similar rates regardless of floral abundance. In contrast, as floral abundance increased, wild pollinator visitation rate decreased more strongly for introduced plants than native plants. Introduced plants as a group and native plants as a group did not differ in bee diversity or preference, but honey bees and wild pollinators preferred different plant species. As a case study, we then focused on knapweed (Centaurea spp.), an introduced plant that was the most preferred plant by honey bees, and that beekeepers value as a late‐summer foraging resource. We compared the extent to which honey bees versus wild pollinators visited knapweed relative to coflowering plants, and we quantified knapweed pollen and nectar collection by honey bees across 22 New York apiaries. Honey bees visited knapweed more frequently than coflowering plants and at a similar rate as all wild pollinators combined. All apiaries contained knapweed pollen in nectar, 86% of apiaries contained knapweed pollen in bee bread, and knapweed was sometimes a main pollen or nectar source for honey bees in late summer. Our results suggest that because of diverging responses to floral abundance and preferences for different plants, honey bees and wild pollinators differ in their use of introduced plants. Depending on the plant and its abundance, removing an introduced plant may impact honey bees more than wild pollinators.