Bee Density Research Articles

Contrasting effects of past and present mass-flowering crop cultivation on bee pollinators shaping yield components in oilseed rape

The cultivation of mass-flowering crops (MFC) can promote pollinators by providing floral resources. However, there is missing knowledge about the effect of MFC cultivation history on bees and their pollination services in agricultural landscapes. We investigated how bee densities in oilseed rape (Brassica napus L.) (OSR) fields were affected by past (cultivation area of the preceding three years) and current MFC area coverages in the surrounding landscape. Moreover, we analyzed how insect pollination, its possible interaction with the plants` pod numbers and MFC covers influence yield components of individual OSR plants and calculated yields (t/ha). To test this, we conducted pollinator surveys and a pollinator exclusion experiment in one oilseed rape field in 17 agricultural landscapes in Germany. We found that wild bee densities were positively affected by past MFC covers and negatively impacted by current OSR covers, indicating enhanced pollinator populations due to previous MFC cultivation and contemporary pollinator dilution. In contrast, honeybees showed opposite responses to past and present MFC cultivation. Furthermore, seed weight per plant of open pollinated plants was positively correlated with past MFC covers. Pollinator exclusion decreased the seed number per pod and increased thousand-seed weight, while yields were unaffected. Pod number interacted with insect pollination in shaping yields, such that pollinator exclusion led to a steeper increase of yield with higher pod numbers. Insect pollination compensated for low pod numbers by increasing the plants` seed number per pod and ultimately yields. Our findings demonstrate a beneficial effect of high MFC covers in the past on bee densities and potentially yield components in the current year. Our study highlights the need for further research on how past and present landscape composition in terms of MFC cultivation interactively affect pollinator communities and their pollination services in agricultural landscapes.

Extreme species density of bees (Apiformes, Hymenoptera) in the warm deserts of North America

Despite the long intertwined evolutionary histories of bees and plants, bee diversity peaks in the xeric areas of the eastern and western hemispheres and not the tropics, where plant diversity is greatest. Intensive sampling in the northeast Chihuahuan Desert of Mexico and the United States provide the first quantitative estimate of bee species richness where high diversity had been predicted in North America from museum records. We find that the density of bee species in a limited area of 16 km2far exceeds any other site in the world and amounts to approximately 14% of the bee species described from the United States. Long-term studies of bees and other pollinators from areas that are minimally impacted by humans provide much-needed baseline data for studies of bees where human impacts are more severe and as climate change accelerates.

Flowering fields, organic farming and edge habitats promote diversity of plants and arthropods on arable land

Abstract Increased farming intensity led to massive declines across multiple farmland taxa. In Europe, measures introduced to counteract these losses include those considered agronomically productive, such as organic farming, as well as those that support no direct production of crops, such as non‐crop flowering fields in conventional farming systems. We studied biodiversity effects of non‐productive flowering fields managed under conventional farming compared to both an organically managed cereal mono‐crop (organic winter spelt fields) and a flowering mixed‐crop (organic lentil mixed‐crop fields) as well as conventionally managed winter wheat fields, which served as control crop. These four crop‐use types were studied on six sites over 3 years (17 sites in total) to assess their impact on the activity density (cover for plants), species richness and community composition of wild plants, carabids, spiders, butterflies and wild bees. Species richness of wild plants was highest under organic farming and at field edges when compared to the interior. In the case of carabids and spiders, species richness was highest at the field edges, but there was no difference between the four crop‐use types. In contrast, activity density and species richness of butterflies and wild bees responded only to flowering crop‐use types, showing no edge effects. Arable land cover in 500 m buffer area also affected community composition of all taxa, with the exception of spiders, but had only minor effects on activity densities and species richness. Synthesis and applications. Our findings underline that there is no single best measure to promote biodiversity on arable land. Instead a mosaic of non‐productive and productive measures such as conventional flowering fields, organic crops and field edge habitats might be more appropriate to support the regional species pool in arable‐dominated landscapes. To support a range of complementary biodiversity‐promoting farming practices, agricultural policy should foster the coordination and collaboration between multiple farmers within the same region by covering additional costs for coordination and prioritizing collaborative schemes.

Higher bee abundance, but not pest abundance, in landscapes with more agriculture on a late-flowering legume crop in tropical smallholder farms.

BackgroundLandscape composition is known to affect both beneficial insect and pest communities on crop fields. Landscape composition therefore can impact ecosystem (dis)services provided by insects to crops. Though landscape effects on ecosystem service providers have been studied in large-scale agriculture in temperate regions, there is a lack of representation of tropical smallholder agriculture within this field of study, especially in sub-Sahara Africa. Legume crops can provide important food security and soil improvement benefits to vulnerable agriculturalists. However, legumes are dependent on pollinating insects, particularly bees (Hymenoptera: Apiformes) for production and are vulnerable to pests. We selected 10 pigeon pea (Fabaceae: Cajunus cajan (L.)) fields in Malawi with varying proportions of semi-natural habitat and agricultural area within a 1 km radius to study: (1) how the proportion of semi-natural habitat and agricultural area affects the abundance and richness of bees and abundance of florivorous blister beetles (Coleoptera: Melloidae), (2) if the proportion of flowers damaged and fruit set difference between open and bagged flowers are correlated with the proportion of semi-natural habitat or agricultural area and (3) if pigeon pea fruit set difference between open and bagged flowers in these landscapes was constrained by pest damage or improved by bee visitation.MethodsWe performed three, ten-minute, 15 m, transects per field to assess blister beetle abundance and bee abundance and richness. Bees were captured and identified to (morpho)species. We assessed the proportion of flowers damaged by beetles during the flowering period. We performed a pollinator and pest exclusion experiment on 15 plants per field to assess whether fruit set was pollinator limited or constrained by pests.ResultsIn our study, bee abundance was higher in areas with proportionally more agricultural area surrounding the fields. This effect was mostly driven by an increase in honeybees. Bee richness and beetle abundances were not affected by landscape characteristics, nor was flower damage or fruit set difference between bagged and open flowers. We did not observe a positive effect of bee density or richness, nor a negative effect of florivory, on fruit set difference.DiscussionIn our study area, pigeon pea flowers relatively late—well into the dry season. This could explain why we observe higher densities of bees in areas dominated by agriculture rather than in areas with more semi-natural habitat where resources for bees during this time of the year are scarce. Therefore, late flowering legumes may be an important food resource for bees during a period of scarcity in the seasonal tropics. The differences in patterns between our study and those conducted in temperate regions highlight the need for landscape-scale studies in areas outside the temperate region.

Coping with the cold and fighting the heat: thermal homeostasis of a superorganism, the honeybee colony

The worldwide distribution of honeybees and their fast propagation to new areas rests on their ability to keep up optimal ‘tropical conditions’ in their brood nest both in the cold and in the heat. Honeybee colonies behave like ‘superorganisms’ where individuals work together to promote reproduction of the colony. Social cooperation has developed strongly in thermal homeostasis, which guarantees a fast and constant development of the brood. We here report on the cooperation of individuals in reaction to environmental variation to achieve thermal constancy of 34–36 °C. The measurement of body temperature together with bee density and in-hive microclimate showed that behaviours for hive heating or cooling are strongly interlaced and differ in their start values. When environmental temperature changes, heat production is adjusted both by regulation of bee density due to migration activity and by the degree of endothermy. Overheating of the brood is prevented by cooling with water droplets and increased fanning, which start already at moderate temperatures where heat production and bee density are still at an increased level. This interlaced change and onset of different thermoregulatory behaviours guarantees a graded adaptation of individual behaviour to stabilise the temperature of the brood.

High honeybee abundances reduce wild bee abundances on flowers in the city of Munich

The increase in managed honeybees (Apis mellifera) in many European cities has unknown effects on the densities of wild bees through competition. To investigate this, we monitored honeybees and non-honeybees from 01 April to 31 July 2019 and 2020 at 29 species of plants representing diverse taxonomic and floral-functional types in a large urban garden in the city of Munich in which the same plant species were cultivated in both years. No bee hives were present in the focal garden, and all bee hives in the adjacent area were closely monitored by interviewing the relevant bee keepers in both 2019 and 2020. Honeybee numbers were similar in April of both years, but increased from May to July 2020 compared to 2019. The higher densities correlated with a significant increase in shifts from wild bee to honeybee visits in May/June/July, while visitor spectra in April 2019 and 2020 remained the same. Most of the species that experienced a shift to honeybee visits in 2020 were visited mostly or exclusively for their nectar. There were no shifts towards increased wild bee visits in any species. These results from a flower-rich garden have implications for the discussion of whether urban bee keeping might negatively impact wild bees. We found clear support that high honeybee densities result in exploitative competition at numerous types of flowers.

Study of the efficiency of creeping clover pollination (Trifolium repens L.) by honey bees (Apis mellifera L.)

The creeping clover (Trifolium repens L.) is a natural allotetraploid allogamous species and is characterized by an entomophilic xenogamous pollination method. One of the reasons for the low yield of clover seeds is the insufficient provision of its flowering grass stands with pollinating insects. The main pollinators are representatives of the order of hymenoptera insects (Himenoptera) from the family of true bees (Apidae) – the honey bee Apis mellifera L. and various species of bumblebees (Bombus). Studies have shown that in the conditions of the Central Non-Chernozem region of Russia, the role of bumblebees in the pollination of creeping clover did not exceed 12-13%. In the presence of bumblebees in different growing seasons in the amount of 30-80 to 110-170 indivi-duals/ha during the mass flowering of creeping clover, they -ensured the formation of seed yields of no more than 90 kg/ha. It is established that honey bees perform up to 87-88% of clover pollination. The highest collections of creeping clover seeds are formed at the flight density of bees of 18-21 thousand individuals per hectare. This makes it possible to form a biological clover yield over 720 kg/ha. To ensure such a rich flight of bees, it is necessary to have at least 2-3 strong bee colonies on clover crops.

Variación en la densidad de poblaciones de Apis mellifera L. en el cultivo de mandarina

The research was carried out in order to determine the variation in population density of Apis mellifera L. along blooming period of Satsuma Owari tangerine (Citrus unshiu Marc.), according to environmental factors at Huaral – Lima. Adults honey bee data were registered on 10 trees, three times a week, on schedules from 8:00 a.m. – 10:00 a.m., 1:00 p.m. – 3:00 p.m. and 3:00 p.m. – 5:00 p.m. on flower buds, opened flowers and withered flowers phases. Besides, aspects of adult bee behavior were registered. Low population values were registered on flower buds and withered flowers phases, while high values were registered on opened flowers phase by the presence of nectar. Higher densities of adult bees were registered at 11:00 a.m. – 1:00 p.m. and 3:00 p.m. – 5:00 p.m. schedules, because of good temperature and solar radiation conditions, while low values were registered on 8:00 a.m. – 10:00 a.m. schedule. The research indicates that tangerine compete against other fruit trees, respect to the visit of bees for harvest of pollen, nectar and pollination, on schedule from 11 a.m. forward on flowering period.

Wildflower plantings on fruit farms provide pollen resources and increase nesting by stem nesting bees

AbstractWildflower plantings on farms have been shown to attract foraging wild bees, however, whether these added floral resources increase nesting densities of bees remains largely untested. We placed nest boxes containing natural reeds at 20 fruit farms in Michigan. We then compared nesting densities between farms with and without wildflower plantings and analysed nest provisions to evaluate use of wildflower plantings for brood provisioning. We found significantly greater nesting at farms with wildflower plantings, with only one out of 236 completed nests at a farm without a planting. The majority of nests were completed by Megachile pugnata, with a portion of nests completed by Osmia caerulescens. We found that nesting bees collected pollen from only a subset of the available flowers in the wildflower plantings, with a strong preference for Centaurea maculosa, and Rudbeckia type pollens. While these species were found growing in the plantings, only Rudbeckia type species were seeded in the plantings. This study provides evidence that wildflower plantings (though not only seeded species) are filling a critical resource gap for stem‐nesting bees in agricultural landscapes and likely support local populations.

Orchard layout and plant traits influence fruit yield more strongly than pollinator behaviour and density in a dioecious crop.

Mutualistic plant-pollinator interactions are critical for the functioning of both non-managed and agricultural systems. Mathematical models of plant-pollinator interactions can help understand key determinants in pollination success. However, most previous models have not addressed pollinator behavior and plant biology combined. Information generated from such a model can inform optimal design of crop orchards and effective utilization of managed pollinators like western honey bees (Apis mellifera), and help generate hypotheses about the effects of management practices and cultivar selection. We expect that the number of honey bees per flower and male to female flower ratio will influence fruit yield. To test the relative importance of these effects, both singly and simultaneously, we utilized a delay differential equation model combined with Latin hypercube sampling for sensitivity analysis. Empirical data obtained from historical records and collected in kiwifruit (Actinidia chinensis) orchards in New Zealand were used to parameterize the model. We found that, at realistic bee densities, the optimal orchard had 65-75% female flowers, and the most benefit was gained from the first 6-8 bees/1000 flowers, with diminishing returns thereafter. While bee density significantly impacted fruit production, plant-based parameters-flower density and male:female flower ratio-were the most influential. The predictive model provides strategies for improving crop management, such as choosing cultivars which have their peak bloom on the same day, increasing the number of flowers with approximately 70% female flowers in the orchard, and placing enough hives to maintain more than 6 bees per 1000 flowers to optimize yield.

Landscape crop diversity and semi-natural habitat affect crop pollinators, pollination benefit and yield

Agricultural intensification has led to the conversion of natural habitats into agricultural fields, increased field sizes and simplified crop rotations. The resulting homogenisation of the landscape has led to a decline in bees, which provide an essential ecosystem service to agriculture. It has been suggested that an increase in landscape crop diversity supports higher biodiversity by providing more diverse and continuous resources without taking land out of agricultural production. We selected 14 faba bean (Vicia faba minor L.) fields in southern Sweden along uncorrelated gradients of landscape crop diversity and proportion of semi-natural habitat within 1.5 km radii surrounding focal fields. Pollinator surveys and pollinator exclusion experiments were conducted to assess whether landscape crop diversity affected pollinator densities, pollinator foraging behaviour (i.e. legitimate flower visitation, nectar robbing or extra-floral nectary visitation), pollination and yield formation. Landscape crop diversity enhanced bumble bee densities. Insect-pollinated faba bean plants produced, on average, 27 % higher bean weight per plant than bagged plants and the insect pollination benefit decreased with increasing semi-natural habitat cover. Bumble bee and honey bee densities, the proportion of nectar robbing bees as well as faba bean yield increased with increasing proportion of semi-natural habitat. Pollinator densities were not the driver of high yields associated with higher proportions of semi-natural habitat because the observed yield increase was unrelated to pollinator densities and driven by bagged plants that were excluded from pollinator visits. Insect pollination, however, clearly decreased the yield gap associated with low proportions of semi-natural habitat in the landscape. Our results highlight that agri-environmental policies should promote the retention of existing semi-natural habitats and encourage landscape crop diversity to provide pollinators with sufficient food and nesting resources.

Floral abundance and bee density affect species-specific foraging patterns of alpine bumble bees

In response to the qualitative and quantitative changes in floral resources, bumble bees flexibly forage multiple plant species throughout the growing season. During the short summer in an alpine ecosystem, the activity of worker bees is maximized in the middle of the season, when the competition for floral resources may be intense. We predicted that the foraging patterns of bumble bees are affected by both relative floral abundance and interactions between bumble bee species. We recorded the floral abundance of individual plant species and the foraging frequency of bumble bees in an alpine meadow during the major flowering period over 3 years. Two bumble bee species were common during the major flowering period. Although they tended to visit abundant floral species, the shorter-tongued species (Bombus hypocrita) showed a more diverse and flexible floral choice than the longer-tongued species (Bombus beaticola). The degree of floral use overlap between two bumble bee species tended to decrease when the foraging density of the longer-tongued species was high. These results indicated that multiple bumble bee species are able to coexist when certain bee species can flexibly change targeting flowers in response to the temporal variations in flowering species and the density of competing bee species. The extent of foraging flexibility is related to the morphological traits of bee species and availability of floral resources.

Population abundance of Varroa destructor and its effects on Apis mellifera scutellata colonies in Kenya.

The devastating effects of Varroa destructor Anderson and Trueman on Western honeybee colonies (Apis mellifera L.) have been well documented. Not only do these mites cause physical damage to parasitized individuals when they feed on them, but they also transmit viruses and other pathogens, weaken colonies and ultimately may cause their death. Unlike the subspecies of European origin, the honeybees of African origin suffer less from mite infestations. Absconding is one of the factors contributing to low V. destructor population in honeybee colonies as it creates a brood-free period. For a long time, researchers hypothesized that absconding was the main mechanism to control the parasite. The effects of V. destructor are well documented under temperate climatic conditions with a break during winter. Therefore, our study aimed at investigating the impact of V. destructor population growth on colony size, absconding and productivity under natural infestation levels of a tropical/subtropical climate with continuous brood production. We measured several characteristics related to the mite populations, the bee colonies and the resources of the bee colonies for a period of 8 months. The seven colonies that absconded during the study period were not influenced by densities of V. destructor. Absconding of the colonies occurred as a result of low numbers of capped brood. Mite densities were generally low throughout the study period (ranged between 26.9 and 59.8 mites per month) but were positively associated with adult bee densities. The amount of open and capped brood was positively associated with densities of V. destructor in the brood and negatively associated with denisities of V. destructor on screen boards, which appeared as extremely important factors that should be monitored regularly alongside colony stores and availability of pollen.

Functional groups of wild bees respond differently to faba bean Vicia faba L. cultivation at landscape scale

Abstract Concerns about insect declines are growing and the provisioning of ecosystem services like pollination may be threatened. To safeguard biodiversity, greening measures were introduced within the reform of the EU's Common Agricultural Policy. One measure commonly applied by farmers is the cultivation of nitrogen fixing crops. Although underlying studies are largely missing, this measure is criticized as providing no significant biodiversity benefit. Using a landscape‐scale approach, we selected 30 paired study landscapes (1 km × 1 km) in Germany, that is, 15 study landscapes with faba bean (FB) fields (Vicia faba L.) and 15 without any grain legumes. Flower‐visiting wild bees were recorded with transect walks at the field margins of different crops using a stratified sampling approach. We analysed the effect of FB cultivation and landscape composition on the abundance and species richness of wild bees as well as on the functional composition of the bee communities. Bumblebee densities (Bombus spp. Latreille) were more than twice as high in FB compared to control landscapes after the flowering of the beans. Non‐Bombus wild bee densities, however, were not affected by FB cultivation, but were enhanced by increasing amounts of semi‐natural habitats (SNH). After the beans' blooming had ceased, FB landscapes had a higher proportion of wild bees collecting pollen from Fabaceae than control landscapes. The community‐weighted means for bee size, measured as intertegular distance, were not affected by FB cultivation, but we found smaller species and species with shorter tongues with an increasing percentage of SNH. Synthesis and applications. The cultivation of faba bean Vicia faba L. strongly increased bumblebee densities throughout the season. This indicates that also on‐field greening measures can support biodiversity. Nevertheless, since only functional groups adapted to faba bean benefit, measures to promote semi‐natural habitats in agricultural landscapes need to be implemented. We conclude that the combination of on‐ and off‐field measures is essential to maintain farmland biodiversity and the Common Agricultural Policy should furthermore promote both.

Analysis of Pollination Services Provided by Wild and Managed Bees (Apoidea) in Wild Blueberry (Vaccinium angustifolium Aiton) Production in Maine, USA, with a Literature Review

Maine is the largest producer of wild blueberry (Vaccinium angustifolium Aiton) in the United States. Pollination comes from combinations of honey bees (Apis mellifera (L.)), commercial bumble bees (Bombus impatiens Cresson), and wild bees. This study addresses (1) previous research addressing wild-blueberry pollination, (2) effects of wild-bee and honey-bee activity densities on fruit set, yield, and crop value, (3) the economic value of wild-bee communities, and (4) economic consequences of pollinator loss. Bee communities were sampled in 40 fields over three years (2010–2012) and bee activity densities were estimated for bumble bees, honey bees, and other wild bees. These data were applied to an economic model to estimate the value of bee taxa. Bumble bees and honey bees predicted fruit set and reduced its spatial heterogeneity. Other wild bees were not significant predictors of fruit set. Yield was predicted by fruit set and field size, but not pest management tactics. Our analysis showed that disruption in supply of honey bees would result in nearly a 30% decrease in crop yield, buffered in part by wild bees that provide “background” levels of pollination. Honey-bee stocking density and, thus, the activity density of honey bees was greater in larger fields, but not for wild bees. Therefore, a decrease in crop yield would be greater than 30% for large fields due to the proportionally greater investment in honey bees in large fields and a relatively lower contribution by wild bees.

Learning of a Mimic Odor within Beehives Improves Pollination Service Efficiency in a Commercial Crop

The growing global demand for pollination services leads producers to consider new strategies in pollinator management to improve its efficiency in agroecosystems [1-3]. Central place foragers, like honeybees, learn floral cues not only in the field but also inside the nest, where resource cues introduced into the hive improve foraging by guiding bees toward the learned stimuli [4]. In this regard, attempts to condition bees with crop-odor-scented food produced ambiguous results and lacked yield measurements [5-7]. To deepen our understanding of the use of odors as part of a precision pollination strategy, we developed a simple synthetic odorant mixture that bees generalized with the natural floral scent of sunflower for hybrid seed production, an economically important and highly pollinator-dependent crop [8]. Encompassing different experimental approaches, our results show that feeding colonies food scented with the sunflower mimic (SM) odor enabled the establishment of olfactory memories that biased bees to the sunflower crop. The offering of a rewarded odor mimicking the sunflower floral fragrance promoted higher foraging activity, increased the proportion of dances advertising the target inflorescences and reduced delays in dance onset, positively affected the density of bees on the crop, and increased yields from 29% to 57% in different sunflower hybrids. This study highlights the role of olfactory learning within the social context of the hive to bias foraging preferences in a novel agricultural environment and suggest that improvements in the tested parameters were due to beeś anticipated response to the sunflower scent.

Assessing chemical control options and their effects on the lesser clover leaf weevil (Hypera nigrirostris) and red clover pollinators in Western Canada

Single-cut red clover (Trifolium pratense L.) is an important forage seed crop in Western Canada due to its winter hardiness and economical value. However, infestation of lesser clover leaf weevil (Hypera nigrirostris F.) can lead to drastic yield losses (over 50%). The aim of this research was to evaluate the effectiveness of insecticides deltamethrin (Decis) and cyantraniliprole (Exirel) in controlling H. nigrirostris in red clover in both field and laboratory conditions and their impacts on the pollinator community. Pest pressure data in red clover fields was tracked during the 2018 and 2019 growing season in six locations. Under the field conditions both deltamethrin and cyantraniliprole were effective at controlling H. nigrirostris numbers. However, only deltamethrin provided rapid pest suppression in 24 h after treatment application. Rearing H. nigrirostris larvae on treated plants in laboratory conditions showed similar results where both insecticides significantly reduced H. nigrirostris numbers in 10–12 days after treatment. The abundances of H. nigrirostris was always negatively related to seed yield regardless of treatment and initial weevil pressure. Seed yield was significantly lower in untreated controls whereas no differences were found between deltamethrin and cyantraniliprole. In contrast, no yield response was found when H. nigrirostris abundance was lower than 4 larvae per 10 shoots. Unexpectedly, densities of bees were not affected by either treatment and were not related to seed yield. Overall, results indicate that both insecticides can provide sufficient H. nigrirostris control, and that pollinator densities are not limiting seed yield, whereas lack of H. nigrirostris management can cause substantial yield losses.

The Ecology of Autogamy in Wild Blueberry (Vaccinium angustifolium Aiton): Does the Early Clone Get the Bee?

Wild blueberry, Vaccinium angustifolium Aiton, for the most part requires cross-pollination. However, there is a continuum across a gradient from zero to 100% in self-compatibility. We previously found by sampling many fields that 20–25% of clones during bloom have high levels of self-compatibility (≥50%). In 2009–2011, and 2015 we studied the ecology of self-pollination in wild blueberry, specifically its phenology and bee recruitment and subsequent bee density on bloom. We found that highly self-compatible clones were predominantly early blooming genotypes in the wild blueberry population. On average, fruit set and berry weight were highest in self-compatible genotypes. The bumble bee community (queens only early in the spring) was characterized by bees that spent large amounts of time foraging in self-compatible plant patches that comprised only a small proportion of the blueberry field, the highest density in the beginning of bloom when most genotypes in bloom were self-compatible. As bloom proceeded in the spring, more plants were in bloom and thus more land area was occupied by blooming plants. The absolute density of bumble bee queens per m2 declined, as a dilution effect, and this probably resulted in lower fruit set throughout the field.

Impact of Invasive Bees on Plant-Pollinator Interactions and Reproductive Success of Plant Species in Mixed Nothofagus antarctica Forests.

Invasive social bees can alter plant-pollinator interactions with detrimental effects on both partners. However, most studies have focused on one invasive bee species, while the interactions among two or more species remain poorly understood. Also, many study sites had a history of invasive bees, being hard to find sites with historical low abundances. In Patagonia, Bombus ruderatus (F.) invasion begun in 1993 and B. terrestris (L.) in 2006. Though honey bees (Apis mellifera L.) introduction started in 1859, their density is still low in some parts. By experimentally increasing honey bee densities, we evaluated the effect of honey bees and bumblebees floral visitation on native pollinator floral visitation, pollen deposition, and reproductive success of three plant species in mixed Nothofagus antarctica forests of northern Patagonia: Oxalis valdiviensis, Mutisia spinosa and Cirsium vulgare. Our results show that exotic bees became the main floral visitors. No negative association was found between invasive bee and native pollinator visitation rates, but there was evidence of potential competition between honey bees and bumblebees. Floral neighborhood diversity played an important role in pollinator behavior. Conspecific pollen deposition was high for all species, while deposition of heterospecific pollen was very high in M. spinosa and C. vulgare. Not as expected, honey bees visitation rate had a negative effect on heterospecific pollen deposition in C. vulgare. For O. valdiviensis, exotic visitation rates increased conspecific pollen deposition, which was positively related to reproductive success. Although exotic bees became main floral visitors, their contribution to reproductive success was only clear for one species.

Performance, Effectiveness, and Efficiency of Honeybees as Pollinators of Coffea arabica (Gentianales, Rubiaceae).

Studies in crop plants analyzing floral biology in conjunction with effectiveness and efficiency of pollinators on pollen transfer and fruit formation are not common, although they are essential to provide better management actions. On this base, we selected a farm in Bahia, Brazil, to study pollination on coffee plants (Coffea arabica L.). Specifically, we want to analyze if nectar traits influence visitor's performance throughout flower lifetime and if honeybees (Apis mellifera scutellata Lepeletier, 1836) are effective and efficient for coffee pollination comparing fertilization and fructification among four experimental treatments: open (OP), wind (WP), cross (HCP), and single-visit bee pollination (SVBP). We found that honeybees collect both nectar and pollen from coffee flowers and transfer pollen on stigmas even after one visit. No differences were found among treatments regarding the number of pollen grains transferred on the stigmas (effectiveness). OP flowers showed a comparative lower efficiency (pollen tubes and fruit set) probably due to pollination failure as those flowers have a higher variability on the number of deposited pollen grains. Two of the treatments (HCP and SVBP) showed higher fertilization (measuring tubes until the end of the style). Pollen loads seem to be limited by a peak of pollen transference by pollinators, followed by the stabilization in the number of pollen grains deposited per stigma. Thus, reproduction of the coffee can be limited by the quality of pollen grains moved by pollinators instead of quantity. Management strategies should focus on monitoring bee density on plants for increasing pollen quality transfer on flowers trough maintaining the adequate proportions of seminatural habitats and/or the number of hives on agricultural fields according to the flowering of the crop.