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Concept: Bees

151

Evidence of inter-species pathogen transmission from managed to wild bees has sparked concern that emerging diseases could be causing or exacerbating wild bee declines. While some pathogens, like RNA viruses, have been found in pollen and wild bees, the threat these viruses pose to wild bees is largely unknown. Here, we tested 169 bees, representing 4 families and 8 genera, for five common honey bee (Apis mellifera) viruses, finding that more than 80% of wild bees harbored at least one virus. We also quantified virus titers in these bees, providing, for the first time, an assessment of viral load in a broad spectrum of wild bees. Although virus detection was very common, virus levels in the wild bees were minimal-similar to or lower than foraging honey bees and substantially lower than honey bees collected from hives. Furthermore, when we experimentally inoculated adults of two different bee species (Megachile rotundata and Colletes inaequalis) with a mixture of common viruses that is lethal to honey bees, we saw no effect on short term survival. Overall, we found that honey bee RNA viruses can be commonly detected at low levels in many wild bee species, but we found no evidence that these pathogens cause elevated short-term mortality effects. However, more work on these viruses is greatly needed to assess effects on additional bee species and life stages.

Concepts: Microbiology, Insect, Honey bee, Beekeeping, Bee, Bumblebee, Pollinators, Bees

78

Neonicotinoid seed dressings have caused concern world-wide. We use large field experiments to assess the effects of neonicotinoid-treated crops on three bee species across three countries (Hungary, Germany, and the United Kingdom). Winter-sown oilseed rape was grown commercially with either seed coatings containing neonicotinoids (clothianidin or thiamethoxam) or no seed treatment (control). For honey bees, we found both negative (Hungary and United Kingdom) and positive (Germany) effects during crop flowering. In Hungary, negative effects on honey bees (associated with clothianidin) persisted over winter and resulted in smaller colonies in the following spring (24% declines). In wild bees (Bombus terrestris and Osmia bicornis), reproduction was negatively correlated with neonicotinoid residues. These findings point to neonicotinoids causing a reduced capacity of bee species to establish new populations in the year following exposure.

Concepts: United Kingdom, Honey bee, Flower, Bee, Bumblebee, Apidae, Pollinators, Bees

69

Bees provide critical pollination services to 87% of angiosperm plants; however, the reliability of these services may become threatened as bee populations decline. Agricultural intensification, resulting in the simplification of environments at the landscape scale, greatly changes the quality and quantity of resources available for female bees to provision their offspring. These changes may alter or constrain the tradeoffs in maternal investment allocation between offspring size, number and sex required to maximize fitness. Here we investigate the relationship between landscape scale agricultural intensification and the size and number of individuals within a wild ground nesting bee species, Andrena nasonii. We show that agricultural intensification at the landscape scale was associated with a reduction in the average size of field collected A. nasonii adults in highly agricultural landscapes but not with the number of individuals collected. Small females carried significantly smaller (40%) pollen loads than large females, which is likely to have consequences for subsequent offspring production and fitness. Thus, landscape simplification is likely to constrain allocation of resources to offspring through a reduction in the overall quantity, quality and distribution of resources.

Concepts: Evolution, Insect, Sex, Flowering plant, Flower, Bee, Resource allocation, Bees

32

It has been suggested that the widespread use of neonicotinoid insecticides threatens bees, but research on this topic has been surrounded by controversy. In order to synthesize which research approaches have been used to examine the effect of neonicotinoids on bees and to identify knowledge gaps, we systematically reviewed research on this subject that was available on the Web of Science and PubMed in June 2015. Most of the 216 primary research studies were conducted in Europe or North America (82%), involved the neonicotinoid imidacloprid (78%), and concerned the western honey bee Apis mellifera (75%). Thus, little seems to be known about neonicotinoids and bees in areas outside Europe and North America. Furthermore, because there is considerable variation in ecological traits among bee taxa, studies on honey bees are not likely to fully predict impacts of neonicotinoids on other species. Studies on crops were dominated by seed-treated maize, oilseed rape (canola) and sunflower, whereas less is known about potential side effects on bees from the use of other application methods on insect pollinated fruit and vegetable crops, or on lawns and ornamental plants. Laboratory approaches were most common, and we suggest that their capability to infer real-world consequences are improved when combined with information from field studies about realistic exposures to neonicotinoids. Studies using field approaches often examined only bee exposure to neonicotinoids and more field studies are needed that measure impacts of exposure. Most studies measured effects on individual bees. We suggest that effects on the individual bee should be linked to both mechanisms at the sub-individual level and also to the consequences for the colony and wider bee populations. As bees are increasingly facing multiple interacting pressures future research needs to clarify the role of neonicotinoids in relative to other drivers of bee declines.

Concepts: Insect, European honey bee, Honey bee, Beekeeping, Bumblebee, Apidae, Pollinators, Bees

27

The Asian giant honeybees (Apis dorsata) build single-comb nests in the open, which makes this species particularly susceptible to environmental strains. Long-term infrared (IR) records documented cool nest regions (CNR) at the bee curtain (nCNR = 207, nnests > 20) distinguished by marked negative gradients (ΔTCNR/d < -3°C / 5 cm) at their margins. CNRs develop and recede within minutes, predominantly at higher ambient temperatures in the early afternoon. The differential size (ΔACNR) and temperature (ΔTCNR) values per time unit correlated mostly positively (RAT > 0) displaying the Venturi effect, which evidences funnel properties of CNRs. The air flows inwards through CNRs, which is verified by the negative spatial gradient ΔTCNR/d, by the positive grading of TCNR with Tamb and lastly by fanners which have directed their abdomens towards CNRs. Rare cases of RAT < 0 (< 3%) document closing processes (for ΔACNR/Δt < -0.4 cm2/s) but also suggest ventilation of the bee curtain (for ΔACNR/Δt > +0.4 cm2/s) displaying “inhalation” and “exhalation” cycling. “Inhalation” could be boosted by bees at the inner curtain layers, which stretch their extremities against the comb enlarging the inner nest lumen and thus causing a pressure fall which drives ambient air inwards through CNR funnels. The relaxing of the formerly “activated” bees could then trigger the “exhalation” process, which brings the bee curtain, passively by gravity, close to the comb again. That way, warm, CO2-enriched nest-borne air is pressed outwards through the leaking mesh of the bee curtain. This ventilation hypothesis is supported by IR imaging and laser vibrometry depicting CNRs in at least four aspects as low-resistance convection funnels for maintaining thermoregulation and restoring fresh air in the nest.

Concepts: Insect, Respiratory physiology, Honey bee, Beekeeping, Bumblebee, Apidae, Bees, Apiology

24

Using precise pollen species determination by conventional microscopic methods, accompanied by molecular genetic markers, we found bees collect GMO (genetically modified) soybean pollen and incorporate it in Yucatan honey. Honey comb samples from Las Flores, Campeche, Mexico, often contained soybean pollen. Pollen in honey was analyzed in nine samples; six contained substantial soy pollen and two tested positive for soybean GMO. Our analyses confirm field observations that honey bees, Apis mellifera, gather soybean pollen and nectar. The resultant risk for honey production in the Yucatán Peninsula and Mexico is evident in wholesale price reduction of 12% when GMO products are detected and honey consignments are rejected. Although this affects only 1% of current export honey (2011-2013) GMO soybean is an unacknowledged threat to apiculture and its economics in one of the world’s foremost honey producing areas.

Concepts: Molecular biology, Insect, Soybean, Honey bee, Beekeeping, Honey, Bees, Honeycomb

17

Eusocial insects live in teeming societies with thousands of their kin. In this crowded environment, workers combat disease by removing or burying their dead or diseased nestmates. For honey bees, we found that hygienic brood-removal behavior is triggered by two odorants - β-ocimene and oleic acid - which are released from brood upon freeze-killing. β-ocimene is a co-opted pheromone that normally signals larval food-begging, whereas oleic acid is a conserved necromone across arthropod taxa. Interestingly, the odorant blend can induce hygienic behavior more consistently than either odorant alone. We suggest that the volatile β-ocimene flags hygienic workers' attention, while oleic acid is the death cue, triggering removal. Bees with high hygienicity detect and remove brood with these odorants faster than bees with low hygienicity, and both molecules are strong ligands for hygienic behavior-associated odorant binding proteins (OBP16 and OBP18). Odorants that induce low levels of hygienic behavior, however, are weak ligands for these OBPs. We are therefore beginning to paint a picture of the molecular mechanism behind this complex behavior, using odorants associated with freeze-killed brood as a model.

Concepts: Death, Insect, Honey bee, Beekeeping, Bee, Ant, Bees, Termite

16

Land-use change and global warming are important factors driving bee decline, but it is largely unknown whether these drivers have resulted in changes in the life-history traits of bees. Recent studies have shown a stronger population decline of large- than small-bodied bee species, suggesting there may have been selective pressure on large, but not on small species to become smaller. Here we test this hypothesis by analyzing trends in bee body size of 18 Dutch species over a 147-year period using specimens from entomological collections. Large-bodied female bees shrank significantly faster than small-bodied female bees (6.5% and 0.5% respectively between 1900 and 2010). Changes in temperature during the flight period of bees did not influence the size-dependent shrinkage of female bees. Male bees did not shrink significantly over the same time period. Our results could imply that under conditions of declining habitat quantity and quality it is advantageous for individuals to be smaller. The size and sex-dependent responses of bees point towards an evolutionary response but genetic studies are required to confirm this. The declining body size of the large bee species that currently dominate flower visitation of both wild plants and insect-pollinated crops may have negative consequences for pollination service delivery.

Concepts: Evolution, Biology, Insect, Honey bee, Flower, Bee, Bumblebee, Bees

16

Early-life social experiences cause lasting changes in behavior and health for a variety of animals including humans, but it is not well understood how social information “gets under the skin” resulting in these effects. Adult honey bees (Apis mellifera) exhibit socially coordinated collective nest defense, providing a model for social modulation of aggressive behavior. Here we report for the first time that a honey bee’s early-life social environment has lasting effects on individual aggression: bees that experienced high-aggression environments during pre-adult stages showed increased aggression when they reached adulthood relative to siblings that experienced low-aggression environments, even though all bees were kept in a common environment during adulthood. Unlike other animals including humans however, high-aggression honey bees were more, rather than less, resilient to immune challenge, assessed as neonicotinoid pesticide susceptibility. Moreover, aggression was negatively correlated with ectoparasitic mite presence. In honey bees, early-life social experience has broad effects, but increased aggression is decoupled from negative health outcomes. Because honey bees and humans share aspects of their physiological response to aggressive social encounters, our findings represent a step towards identifying ways to improve individual resiliency. Pre-adult social experience may be crucial to the health of the ecologically threatened honey bee.

Concepts: Insect, Honey bee, Beekeeping, Bumblebee, Apidae, Bees, Apiology, Bees and toxic chemicals

12

Neonicotinoid insecticides have been identified as an important factor contributing to bee diversity declines. Nonetheless, uncertainties remain about their impact under field conditions. Most studies have been conducted on Apis mellifera and tested single compounds. However, in agricultural environments, bees are often exposed to multiple pesticides. We explore synergistic mortality between a neonicotinoid (clothianidin) and an ergosterol-biosynthesis-inhibitor fungicide (propiconazole) in three bee species (A. mellifera, Bombus terrestris, Osmia bicornis) following oral exposure in the laboratory.

Concepts: Insect, Honey bee, Bee, Bumblebee, Insecticide, Apidae, Pollinators, Bees