The habitat disruption induces immune-suppression and oxidative stress in honey bees

The honey bee is a major insect used for pollination of many commercial crops worldwide. Although the use of honey bees for pollination can disrupt the habitat, the effects on their physiology have never been determined. Recently, honey bee colonies have often collapsed when introduced in greenhouses for pollination in Japan. Thus, suppressing colony collapses and maintaining the number of worker bees in the colonies is essential for successful long-term pollination in greenhouses and recycling of honey bee colonies. To understand the physiological states of honey bees used for long-term pollination in greenhouses, we characterized their gene expression profiles by microarray. We found that the greenhouse environment changes the gene expression profiles and induces immune-suppression and oxidative stress in honey bees. In fact, the increase of the number of Nosema microsporidia and protein carbonyl content was observed in honey bees during pollination in greenhouses. Thus, honey bee colonies are likely to collapse during pollination in greenhouses when heavily infested with pathogens. Degradation of honey bee habitat by changing the outside environment of the colony, during pollination services for example, imposes negative impacts on honey bees. Thus, worldwide use of honey bees for crop pollination in general could be one of reasons for the decline of managed honey bee colonies.     

Differential sensitivity of honey bees and bumble bees to a dietary insecticide (imidacloprid)

Currently, there is concern about declining bee populations and the sustainability of pollination services. One potential threat to bees is the unintended impact of systemic insecticides, which are ingested by bees in the nectar and pollen from flowers of treated crops. To establish whether imidacloprid, a systemic neonicotinoid and insect neurotoxin, harms individual bees when ingested at environmentally realistic levels, we exposed adult worker bumble bees, Bombus terrestris L. (Hymenoptera: Apidae), and honey bees, Apis mellifera L. (Hymenoptera: Apidae), to dietary imidacloprid in feeder syrup at dosages between 0.08 and 125 μg l^?1. Honey bees showed no response to dietary imidacloprid on any variable that we measured (feeding, locomotion and longevity). In contrast, bumble bees progressively developed over time a dose-dependent reduction in feeding rate with declines of 10–30% in the environmentally relevant range of up to 10 μg l^?1, but neither their locomotory activity nor longevity varied with diet. To explain their differential sensitivity, we speculate that honey bees are better pre-adapted than bumble bees to feed on nectars containing synthetic alkaloids, such as imidacloprid, by virtue of their ancestral adaptation to tropical nectars in which natural alkaloids are prevalent. We emphasise that our study does not suggest that honey bee colonies are invulnerable to dietary imidacloprid under field conditions, but our findings do raise new concern about the impact of agricultural neonicotinoids on wild bumble bee populations.     

Native bees provide insurance against ongoing honey bee losses

One of the values of biodiversity is that it may provide 'biological insurance' for services currently rendered by domesticated species or technology. We used crop pollination as a model system, and investigated whether the loss of a domesticated pollinator (the honey bee) could be compensated for by native, wild bee species. We measured pollination provided to watermelon crops at 23 farms in New Jersey and Pennsylvania, USA, and used a simulation model to separate the pollen provided by honey bees and native bees. Simulation results predict that native bees alone provide sufficient pollination at > 90% of the farms studied. Furthermore, empirical total pollen deposition at flowers was strongly, significantly correlated with native bee visitation but not with honey bee visitation. The honey bee is currently undergoing extensive die-offs because of Colony Collapse Disorder. We predict that in our region native bees will buffer potential declines in agricultural production because of honey bee losses.     

Influence of brood, vent screening, and time of year on honey bee (Hymenoptera: Apidae) pollination and fruit quality of greenhouse tomatoes

Greenhouse tomatoes, Lycopersicon esculentum Miller (Solanaceae), are autogamous, but facilitated pollination results in increased fruit size and set. Previous research examining honey bee pollination in greenhouse tomato crops established that fruit quality resulting from honey bee visitation is often comparable to bumble bees (Bombus spp.) and significantly better than in flowers that receive no facilitated pollination. However, management alternatives have not been studied to improve tomato fruit quality when honey bees are the only pollination option available for the high-value greenhouse industry. We investigated whether the quantity of brood (eggs, larvae, and pupae) in a honey bee colony in the winter and screening on greenhouse vents in the summer would encourage honey bee foraging on tomato flowers. We also established the influence of time of year on the potential for honey bees to be effective pollinating agents. We constructed small honey bee colonies full of naive forager bees with either two frames of brood ("brood colonies") or two empty frames ("no-brood") and compared total fruit set and the number of tomato seeds resulting from fruit potentially visited by honey bees in each of these treatments to bagged flowers that received no facilitated pollination. There was no significant difference in the quality of fruit resulting from honey bees from "brood" and "no-brood" colonies. However, these fruits produced significantly more seeds than bagged flowers restricted from facilitated pollination. Honey bees from brood and no-brood colonies also resulted in 98% fruit set compared with 80% fruit set in bagged flowers that received no facilitated pollination. During the summer, the number of seeds per fruit did not differ significantly between unbagged flowers potentially visited by honey bees in screened greenhouses and unscreened greenhouses and bagged flowers that received no facilitated pollination. However, time of year did have a significant influence on the quality of fruit produced by honey bees compared with flowers that received no facilitated pollination, because no difference in seed number was observed between the treatments after mid-April. The results from this study demonstrate that the management of brood levels and vent screening cannot be used to improve the quality of fruit resulting from honey bee pollination and that honey bees can be a feasible greenhouse pollination alternative only during the winter.     

Imidacloprid Alters Foraging and Decreases Bee Avoidance of Predators

Concern is growing over the effects of neonicotinoid pesticides, which can impair honey bee cognition. We provide the first demonstration that sublethal concentrations of imidacloprid can harm honey bee decision-making about danger by significantly increasing the probability of a bee visiting a dangerous food source. Apis cerana is a native bee that is an important pollinator of agricultural crops and native plants in Asia. When foraging on nectar containing 40 µg/L (34 ppb) imidacloprid, honey bees (Apis cerana) showed no aversion to a feeder with a hornet predator, and 1.8 fold more bees chose the dangerous feeder as compared to control bees. Control bees exhibited significant predator avoidance. We also give the first evidence that foraging by A. cerana workers can be inhibited by sublethal concentrations of the pesticide, imidacloprid, which is widely used in Asia. Compared to bees collecting uncontaminated nectar, 23% fewer foragers returned to collect the nectar with 40 µg/L imidacloprid. Bees that did return respectively collected 46% and 63% less nectar containing 20 µg/L and 40 µg/L imidacloprid. These results suggest that the effects of neonicotinoids on honey bee decision-making and other advanced cognitive functions should be explored. Moreover, research should extend beyond the classic model, the European honey bee (A. mellifera), to other important bee species.     

Long‐term risk assessment on noneffective and effective toxic doses of imidacloprid to honeybee workers

Imidacloprid is the most widely used insecticide in agriculture. Its impact on honeybees has received worldwide attention. Foliar sprays are commonly and frequently used for piercing insect control, particularly on cotton in southern USA. To simulate field exposures of formulated imidacloprid (Advise^® 2Fl), we used a modified spray tower to treat honeybee workers and monitored five enzyme activities and survival for up to 52 days. Results indicated that spray treatments twice a week for 52 days with 0.001 and 1 mg/L and once a week for three weeks with 4.3 mg/L Advise showed no adverse effect on bee survival, where imidacloprid‐treated bees could live as long as untreated bees. Concentration ≥80 mg/L significantly reduced bee survival, and substantial number of bees continued to die after 48‐hr of post‐treatment period which was commonly used for measuring insecticide toxicity. The body weight of imidacloprid‐treated bees (at LC₂₀ and LC₅₀) was also significantly reduced. Enzymatic data showed that activities of detoxification enzymes esterase and glutathione S‐transferase (GST), insecticide‐target enzyme acetylcholinesterase (AChE) and honey enzyme invertase in imidacloprid‐treated survivors were mostly similar to those found in untreated bees. The immunity‐related phenoloxidase (PO) activity in imidacloprid‐treated survivors was also mostly similar to that of untreated control, but higher PO activity was detected in bees treated with higher concentrations for 3 weeks. By using both bioassays and enzymatic assays, this study revealed long‐term noneffective and effective concentrations of imidacloprid that may be useful for accurate assessment of toxicity risk of neonicotinoids to bees.     

Bumble bee abundance and richness improves honey bee pollination behaviour in sweet cherry

Pollination service in agricultural crops increases significantly with pollinator diversity and wild pollinator abundance. Differences in the foraging behaviour of pollinating insects are one of the reasons why pollinator diversity and abundance enhances crop pollination. Here, we focused on the foraging behaviour of honey bees and bumble bees in sweet cherry orchards. In addition, we studied the influence of bee diversity and abundance on the foraging behaviour of honey bees and bumble bees. Honey bees were found to visit fewer flowers than bumble bees. Bumble bees also showed a higher probability of changing trees between rows than honey bees. Both visitation rate and probability of row changes of honey bees increased with bumble bee diversity and with bumble bee abundance. We also found that the probability of row changes of honey bees increased with increasing bumble bee abundance. These effects of bumble bee richness and abundance on the pollination behaviour of honey bees can improve the pollination performance of honey bees in crops that depend on cross pollination. Our results highlight the higher pollination performance of bumble bees and the facilitative effect of wild pollinators to crop pollination.     

Honey bee (Hymenoptera: Apidae) distribution and potential for supplementary pollination in commercial tomato greenhouses during winter

This study examined the use of honey bees, Apis mellifera L., to supplement bumble bee, Bombus spp., pollination in commercial tomato, Lycopersicon esculentum Miller, greenhouses in Western Canada. Honey bee colonies were brought into greenhouses already containing bumble bees and left for 1 wk to acclimatize. The following week, counts of honey and bumble bees foraging and flying throughout the greenhouse were conducted three times per day, and tomato flowers open during honey bee pollination were marked for later fruit harvest. The same counts and flower-marking also were done before and after the presence of honey bees to determine the background level of bumble bee pollination. Overall, tomato size was not affected by the addition of honey bees, but in one greenhouse significantly larger tomatoes were produced with honey bees present compared with bumble bees alone. In that greenhouse, honey bee foraging was greater than in the other greenhouses. Honey bees generally foraged within 100 m of their colony in all greenhouses. Our study invites further research to examine the use of honey bees with reduced levels of bumble bees, or as sole pollinators of greenhouse tomatoes. We also make specific recommendations for how honey bees can best be managed in greenhouses.     

Pollination Services Provided by Bees in Pumpkin Fields Supplemented with Either Apis mellifera or Bombus impatiens or Not Supplemented

Pollinators provide an important service in many crops. Managed honey bees (Apis mellifera L.) are used to supplement pollination services provided by wild bees with the assumption that they will enhance pollination, fruit set and crop yield beyond the levels provided by the wild bees. Recent declines in managed honey bee populations have stimulated interest in finding alternative managed pollinators to service crops. In the eastern U.S., managed hives of the native common eastern bumble bee (Bombus impatiens Cresson) may be an excellent choice. To examine this issue, a comprehensive 2-yr study was conducted to compare fruit yield and bee visits to flowers in pumpkin (Cucurbita pepo L.) fields that were either supplemented with A. mellifera hives, B. impatiens hives or were not supplemented. We compared pumpkin yield, A. mellifera flower visitation frequency and B. impatiens flower visitation frequency between treatments. Results indicated that supplementing pumpkin fields with either A. mellifera or B. impatiens hives did not increase their visitation to pumpkin flowers or fruit yield compared with those that were not supplemented. Next, the relationship between frequency of pumpkin flower visitation by the most prominent bee species (Peponapis pruinosa (Say), B. impatiens and A. mellifera) and fruit yield was determined across all pumpkin fields sampled. Fruit yield increased as the frequency of flower visits by A. mellifera and B. impatiens increased in 2011 and 2012, respectively. These results suggest that supplementation with managed bees may not improve pumpkin production and that A. mellifera and B. impatiens are important pollinators of pumpkin in our system.     

生态条件的多样性变化对蜜蜂生存的影响

蜜蜂在整个生态系统中起着重要的传花授粉作用,是生态链中不可或缺的物种。随着现代农业的发展,蜜蜂赖以生存的环境遭到破坏,继而引发蜜蜂数量大幅减少,影响了蜂种的生存与可持续发展。总结了近年来生态条件的变化,归纳了影响蜜蜂生存的主要因素,分析了蜜蜂生存艰难的原因,提出了蜜蜂生存的关键问题,并展望了未来维持蜜蜂强群的主要研究方向。     

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.     

Seed set in Guayule (Parthenium argentatum,asteraceae) in relation to insect pollination

Guayule (Tarthenium argentatum, Asteraceae) is one of two major plant species grown for natural rubber. Studies were conducted to determine the effect of honey bee (Apis mellifera) pollination and season on seed set and total seed yield/ha. The experiments involved four pollination treatments: plants caged with bees; plants caged without bees; plants open pollinated (uncovered); and plants individually covered. Seeds were harvested monthly July–September 1984, and May–September 1985. Plots with bees produced at least 150% more seeds than plots without bees, and there were no qualitative differences in the seed weights among treatments. Highest seed yield was in May and September. Results indicate that (1) insect pollination in guayule increases seed yield and (2) fewer seeds are produced in the warmest months.     

Farming practices influence wild pollinator populations on squash and pumpkin

Recent declines in managed honey bee, Apis mellifera L., colonies have increased interest in the current and potential contribution of wild bee populations to the pollination of agricultural crops. Because wild bees often live in agricultural fields, their population density and contribution to crop pollination may be influenced by farming practices, especially those used to reduce the populations of other insects. We took a census of pollinators of squash and pumpkin at 25 farms in Virginia, West Virginia, and Maryland to see whether pollinator abundance was related to farming practices. The main pollinators were Peponapis pruinosa Say; honey bees, and bumble bees (Bombus spp.). The squash bee was the most abundant pollinator on squash and pumpkin, occurring at 23 of 25 farms in population densities that were commonly several times higher than that of other pollinators. Squash bee density was related to tillage practices: no-tillage farms hosted three times as great a density of squash bees as tilled farms. Pollinator density was not related to pesticide use. Honey bee density on squash and pumpkin was not related to the presence of managed honey bee colonies on farms. Farms with colonies did not have more honey bees per flower than farms that did not keep honey bees, probably reflecting the lack of affinity of honey bees for these crops. Future research should examine the economic impacts of managing farms in ways that promote pollinators, particularly pollinators of crops that are not well served by managed honey bee colonies.     

Honey bee (Hymenoptera: Apidae) foraging in response to preconditioning with onion flower scent compounds

Onion (Allium cepa L.) seed production has long been plagued with yield problems because of lack of pollination by the honey bee, Apis mellifera L. To attempt to attract more pollinators to the onion seed production field, honey bees were conditioned to associate onion floral odor components with a reward. Isolated nucleus hives of honey bees were fed 30% sucrose solutions scented with a 0.2% solution of onion floral odor compounds. After feeding on these solutions for 6 wk, bees were not found to prefer onion flowers to two competing food sources, carrot and alfalfa flowers, at the 5% significance level. However, there was an overall trend indicating a change in honey bee behavior, with fewer "trained" bees visiting alfalfa and carrot and more visiting onion. Thus, it may be possible to alter honey bee behavior with preconditioning but probably not to a degree that would be economically significant.     

Orchard pollination in Capitol Reef National Park,Utah, USA. Honey bees or native bees?

Capitol Reef National Park in central Utah, USA surrounds 22 managed fruit orchards started over a century ago by Mormon pioneers. Honey bees are imported for pollination, although the area in which the Park is embedded has over 700 species of native bees, many of which are potential orchard pollinators. We studied the visitation of native bees to apple, pear, apricot, and sweet cherry over 2 years. Thirty species of bees visited the flowers but, except for pear flowers, most were uncommon compared to honey bees. Evidence that honey bees prevented native bees from foraging on orchard crop flowers was equivocal: generally, honey bee and native bee visitation rates to the flowers were not negatively correlated, nor were native bee visitation rates positively correlated with distance of orchards from honey bee hives. Conversely, competition was tentatively suggested by much larger numbers of honey bees than natives on the flowers of apples, apricots and cherry; and by the large increase of native bees on pears, where honey bee numbers were low. At least one-third of the native bee species visiting the flowers are potential pollinators, including cavity-nesting species such as Osmia lignaria propinqua, currently managed for small orchard pollination in the US, plus several fossorial species, including one rosaceous flower specialist (Andrena milwaukiensis). We suggest that gradual withdrawal of honey bees from the Park would help conserve native bee populations without decreasing orchard crop productivity, and would serve as a demonstration of the commercial value of native pollinators.     

Managing honey bees (Hymenoptera: Apidae) for greenhouse tomato pollination

Although commercially reared colonies of bumble bees (Bombus sp.) are the primary pollinator world-wide for greenhouse tomatoes (Lycopersicon esculentum Mill.) previous research indicates that honey bees (Apis mellifera L.) might be a feasible alternative or supplement to bumble bee pollination. However, management methods for honey bee greenhouse tomato pollination scarcely have been explored. We 1) tested the effect of initial amounts of brood on colony population size and flight activity in screened greenhouses during the winter, and 2) compared foraging from colonies with brood used within screened and unscreened greenhouses during the summer. Brood rearing was maintained at low levels in both brood and no-brood colonies after 21 d during the winter, and emerging honey bees from both treatments had significantly lower weights than bees from outdoor colonies. Honey bee flight activity throughout the day and over the 21 d in the greenhouse was not influenced by initial brood level. In our summer experiment, brood production in screened greenhouses neared zero after 21 d but higher levels of brood were reared in unscreened greenhouses with access to outside forage. Flower visitation measured throughout the day and over the 21 d the colonies were in the greenhouse was not influenced by screening treatment. An economic analysis indicated that managing honey bees for greenhouse tomato pollination would be financially viable for both beekeepers and growers. We conclude that honey bees can be successfully managed for greenhouse tomato pollination in both screened and unscreened greenhouses if the foraging force is maintained by replacing colonies every 3 wk.     

Management increases genetic diversity of honey bees via admixture

The process of domestication often brings about profound changes in levels of genetic variation in animals and plants. The honey bee, Apis mellifera, has been managed by humans for centuries for both honey and wax production and crop pollination. Human management and selective breeding are believed to have caused reductions in genetic diversity in honey bee populations, thereby contributing to the global declines threatening this ecologically and economically important insect. However, previous studies supporting this claim mostly relied on population genetic comparisons of European and African (or Africanized) honey bee races; such conclusions require reassessment given recent evidence demonstrating that the honey bee originated in Africa and colonized Europe via two independent expansions. We sampled honey bee workers from two managed populations in North America and Europe as well as several old-world progenitor populations in Africa, East and West Europe. Managed bees had highly introgressed genomes representing admixture between East and West European progenitor populations. We found that managed honey bees actually have higher levels of genetic diversity compared with their progenitors in East and West Europe, providing an unusual example whereby human management increases genetic diversity by promoting admixture. The relationship between genetic diversity and honey bee declines is tenuous given that managed bees have more genetic diversity than their progenitors and many viable domesticated animals.     

The Potential Influence of Bumble Bee Visitation on Foraging Behaviors and Assemblages of Honey Bees on Squash Flowers in Highland Agricultural Ecosystems

Bee species interactions can benefit plant pollination through synergistic effects and complementary effects, or can be of detriment to plant pollination through competition effects by reducing visitation by effective pollinators. Since specific bee interactions influence the foraging performance of bees on flowers, they also act as drivers to regulate the assemblage of flower visitors. We selected squash (Cucurbita pepo L.) and its pollinators as a model system to study the foraging response of honey bees to the occurrence of bumble bees at two types of sites surrounded by a high amount of natural habitats (≥ 58% of land cover) and a low amount of natural habitats (≤ 12% of land cover) in a highland agricultural ecosystem in China. At the individual level, we measured the elapsed time from the departure of prior pollinator(s) to the arrival of another pollinator, the selection of honey bees for flowers occupied by bumble bees, and the length of time used by honey bees to explore floral resources at the two types of sites. At the community level, we explored the effect of bumble bee visitation on the distribution patterns of honey bees on squash flowers. Conclusively, bumble bee visitation caused an increase in elapsed time before flowers were visited again by a honey bee, a behavioral avoidance by a newly-arriving honey bee to select flowers occupied by bumble bees, and a shortened length of time the honey bee takes to examine and collect floral resources. The number of overall bumble bees on squash flowers was the most important factor explaining the difference in the distribution patterns of honey bees at the community level. Furthermore, decline in the number of overall bumble bees on the squash flowers resulted in an increase in the number of overall honey bees. Therefore, our study suggests that bee interactions provide an opportunity to enhance the resilience of ecosystem pollination services against the decline in pollinator diversity.     

Managed bumble bees increase flower visitation but not fruit weight in polytunnel strawberry crops

Animal-mediated pollination is essential for the production and quality of fruits and seeds of many crops consumed by humans. However, crop pollination services might be compromised when wild pollinators are scarce. Managed pollinators are commonly used in crops to supplement such services with the assumption that they will enhance crop yield. However, information on the spatiotemporal pollinator-dependence of crops is still limited. We assessed the contribution of commercial bumble bee colonies compared to the available pollinator community on strawberry (‘Fortuna’ variety) flower visitation and strawberry quality across a landscape gradient of agricultural intensification (i.e. polytunnel berry crop cover). We used colonies of bumble bees in winter and in spring, i.e. when few and most wild pollinators are in their flight period, respectively. The placement of colonies increased visits of bumble bees to strawberry flowers, especially in winter. The use of bumble bee colonies did not affect flower visitation by other insects, mainly honey bees, hoverflies and other Diptera. Flower visitation by both honey bees and wild insects did not vary between seasons and was unrelated to the landscape gradient of berry crop cover. Strawberries were of the highest quality (i.e. weight) when insect-mediated pollination was allowed, and their quality was positively related to wild flower visitors in winter but not in spring. However, increased visits to strawberry flowers by managed bumble bees and honey bees had no effect on strawberry weight. Our results suggest that the pollination services producing high quality strawberry fruits are provided by the flower visitor community present in the study region without the need to use managed bumble bees.     

Key factors influencing forager distribution across macadamia orchards differ among species of managed bees

To achieve maximised and sustainable crop productivity, it is critical that we develop crop-specific strategies for managing pollination. Honey bees (Apis mellifera) and stingless bees (Tetragonula carbonaria) are considered effective pollinators of macadamia (Macadamia integrifolia). The introduction of managed honey bee or stingless bee hives into orchards is likely to boost the numbers of these insects visiting flowers; however, there is a lack of published information and consensus regarding their management for pollination. Here, we identify factors that affect the distribution of both honey bees and stingless bees across cultivated macadamia, and establish whether increased flower visitation leads to higher nut set. A gradient of bee visitation rates was created by placing colonies on the ends of a four-hectare block, and mixed-effect models were applied to assess forager abundance and nut set with respect to distance from hive, time of day, cultivar, and floral display size. Distance from colony had a strong effect on stingless bee numbers, with >96% of individuals recorded within 100 metres of colonies, whereas the distribution of honey bees was more closely related to daily floral display: trees with greater numbers of flowers attracted more honey bees. Simplified surveys conducted in a further 17 macadamia blocks confirm that these are broadly occurring distribution patterns. Bee abundance alone did not significantly predict nut production; however, an indirect effect of bee visits to flowers is inferred, as nut production increased with size of floral display. To encourage a more even distribution of bees and uniform pollination, we recommend placement of stingless bee hives to maximise their distribution through a block (e.g. at 100-m intervals) and management practices that promote even distributions of flowers across trees.