Functionality of Varroa-resistant honey bees (Hymenoptera: Apidae) when used in migratory beekeeping for crop pollination

Two types of honey bees, Apis mellifera L. (Hymenoptera: Apidae), bred for resistance to Varroa destructor Anderson & Trueman were evaluated for performance when used in migratory crop pollination. Colonies of Russian honey bees (RHB) and outcrossed bees with Varroa-sensitive hygiene (VSH) were managed without miticide treatments and compared with colonies of Italian honey bees that served as controls. Control colonies were managed as groups which either were treated twice each year against V. destructor (CT) or kept untreated (CU). Totals of 240 and 247 colonies were established initially for trials in 2008 and 2009, respectively. RHB and VSH colonies generally had adult and brood populations similar to those of the standard CT group regarding pollination requirements. For pollination of almonds [Prunus dulcis (Mill.) D.A.Webb] in February, percentages of colonies meeting the required six or more frames of adult bees were 57% (VSH), 56% (CT), 39% (RHB), and 34% (CU). RHB are known to have small colonies in early spring, but this can be overcome with appropriate feeding. For later pollination requirements in May to July, 94-100% of colonies in the four groups met pollination size requirements for apples (Malus domestica Borkh.), cranberries (Vaccinium macrocarpon Aiton), and lowbush blueberries (Vaccinium angustifolium Aiton). Infestations with V. destructor usually were lowest in CT colonies and tended to be lower in VSH colonies than in RHB and CU colonies. This study demonstrates that bees with the VSH trait and pure RHB offer alternatives for beekeepers to use for commercial crop pollination while reducing reliance on miticides. The high frequency of queen loss (only approximately one fourth of original queens survived each year) suggests that frequent requeening is necessary to maintain desired genetics.     

Effects of fluvalinate and coumaphos on queen honey bees (Hymenoptera: Apidae) in two commercial queen rearing operations

We conducted research on the potential impacts of fluvalinate and coumaphos on honey bee, Apis mellifera L., queen viability and health. Queens were reared in colonies that had been treated with differing amounts of both fluvalinate and coumaphos. Pre- and posttreatment samples of both wax and bees were collected from all of the colonies and analyzed for total concentrations of fluvalinate and coumaphos. All queens were measured for queen weight, ovarial weight, and number of sperm in the spermathecae. The queens treated with high doses of fluvalinate weighed significantly less than low-dose or control queens, but otherwise appeared to develop normally. The highest fluvalinate concentrations were observed in the wax and queen cells of the high-dose group. The developing queens in colonies treated with as little as one coumaphos-impregnated strip for more than 24 h suffered a high mortality rate. Several of the queens showed sublethal effects from the coumaphos, including physical abnormalities and atypical behavior. The queens exposed to coumaphos weighed significantly less and had lower ovary weights than the control group queens. The highest coumaphos concentrations were observed in the queen cells and wax of the high-dose groups.     

American foulbrood and African honey bees (Hymenoptera: Apidae)

We have taken samples of honey from individual beekeepers (N = 64), and of domestic (N = 35) and imported honey (N = 15) retailed in supermarkets in several sub-Saharan countries and cultivated these samples for Paenibacillus larvae subsp. larvae Heyndrickx et al. causing American foulbrood in honey bee colonies. The results are compared with samples of similar backgrounds and treated the same way but collected in Sweden (N = 35). No P. larvae subsp. larvae spores were found in any honey produced in Africa south of the Sahara although honey imported into this region frequently contains the pathogen. Swedish honey frequently contains P. larvae subsp. larvae spores although the general level of visibly infected bee colonies is low (roughly 0.5%). The results suggest that large parts of Africa may be free from American foulbrood. Behavioral studies (hygienic behavior) on Apis mellifera subsp. scutellata Lepeletier in Zimbabwe suggest that hygienic behavior of African bees could influence the apparent low level, or even absence of American foulbrood in large parts of Africa.     

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.     

Resistance to Varroa destructor (Mesostigmata: Varroidae) when mite-resistant queen honey bees (Hymenoptera: Apidae) were free-mated with unselected drones.

This study demonstrated (1) that honey bees, Apis mellifera L, can express a high level of resistance to Varroa destructor Anderson & Trueman when bees were selected for only one resistant trait (suppression of mite reproduction); and (2) that a significant level of mite-resistance was retained when these queens were free-mated with unselected drones. The test compared the growth of mite populations in colonies of bees that each received one of the following queens: (1) resistant--queens selected for suppression of mite reproduction and artificially inseminated in Baton Rouge with drones from similarly selected stocks; (2) resistant x control--resistant queens, as above, produced and free-mated to unselected drones by one of four commercial queen producers; and (3) control--commercial queens chosen by the same four queen producers and free-mated as above. All colonies started the test with approximately 0.9 kg of bees that were naturally infested with approximately 650 mites. Colonies with resistant x control queens ended the 115-d test period with significantly fewer mites than did colonies with control queens. This suggests that beekeepers can derive immediate benefit from mite-resistant queens that have been free-mated to unselected drones. Moreover, the production and distribution of these free-mated queens from many commercial sources may be an effective way to insert beneficial genes into our commercial population of honey bees without losing the genetic diversity and the useful beekeeping characteristics of this population.     

Brood pheromone regulates foraging activity of honey bees (Hymenoptera: Apidae)

Brood pheromone modulated the foraging behavior of commercial honey bee, Apis mellifera L., colonies pollinating a 10-ha market garden of cucumber, Cucurbita pepo L., and zucchini, Cucumis saticus L., in Texas in late autumn. Six colonies were randomly selected to receive 2000 larval equivalents of brood pheromone and six received a blank control. The ratio of pollen to nonpollen foragers entering colonies was significantly greater in pheromone-treated colonies 1 h after treatment. Pheromone-treated foragers returned with pollen load weights that were significantly heavier than controls. Pollen returned by pheromone-treated foragers was 43% more likely to originate from the target crop. Number of pollen grains washed from the bodies of nonpollen foragers from pheromone-treated colonies was significantly greater than controls and the pollen was 54% more likely to originate from the target crop. Increasing the foraging stimulus environment with brood pheromone increased colony-level foraging and individual forager efforts. Brood pheromone is a promising technology for increasing the pollination activity and efficiency of commercial honey bee colonies.     

Nonrandom visitation of brood cells by worker honey bees (Hymenoptera: Apidae)

The visitation pattern by worker honey bees to cells in the brood nest was monitored on an artificially created brood pattern consisting of about one-fourth brood cells evenly distributed among empty cells. The majority (63 %) of the observed workers selectively entered larval cells. In contrast, some workers avoided egg cells when presented a choice of egg vs empty cells. The results suggest that larvae produce a general signal indicating their presence to worker bees. Eggs also seem to produce a signal, which is perceived to be different from the one from larvae.     

Application of a modified selection index for honey bees (Hymenoptera: Apidae)

Nine different genetic families of honey bees (Apis mellifera L.) were compared using summed z-scores (phenotypic values) and a modified selection index (Imod). Imod values incorporated both the phenotypic scores of the different traits and the economic weightings of these traits, as determined by a survey of commercial Ontario beekeepers. Largely because of the high weight all beekeepers place on honey production, a distinct difference between line rankings based on phenotypic scores and Imod scores was apparent, thereby emphasizing the need to properly weight the traits being evaluated to select bee stocks most valuable for beekeepers. Furthermore, when beekeepers who made >10% of their income from queen and nucleus colony sales assigned relative values to the traits used in the Imod calculations, the results differed from those based on weightings assigned by honey producers. Our results underscore the difficulties the North American beekeeping industry must overcome to devise effective methods of evaluating colonies for breeding purposes.     

Africanized honey bees (Hymenoptera: Apidae) have a greater fidelity to sunflowers than European bees

A study of sunflower, Helianthus annuus L., pollen collection by Africanized and European honey bees, Apis mellifera L., was conducted in a hybrid seed production field in Argentina. Africanized honey bees collected significantly larger proportions of sunflower pollen than did European honey bees. The result suggests that Africanized bees would be more efficient for commercial sunflower seed production.     

Alarm pheromone perception in honey bees is decreased by smoke (Hymenoptera: Apidae)

The application of smoke to honey bee(Apis mellifera) antennae reduced the subsequent electroantennograph response of the antennae to honey bee alarm pheromones, isopentyl acetate, and 2-heptanone. This effect was reversible, and the responsiveness of antennae gradually returned to that of controls within 10–20 min. A similar effect occurred with a floral odor, phenylacetaldehyde, suggesting that smoke interferes with olfaction generally, rather than specifically with honey bee alarm pheromones. A reduction in peripheral sensitivity appears to be one component of the mechanism by which smoke reduces nest defense behavior of honey bees.     

Environmental influences on flight activity of USDA-ARS Russian and Italian stocks of honey bees (Hymenoptera: Apidae) during almond pollination

Differences in flight activity and in the percentages of pollen foragers between commercially managed honey bees, Apis mellifera L. (Hymenoptera: Apidae), of two stocks (USDA-ARS Russian, n = 41 colonies; and Italian, n = 43 colonies) were evaluated in an almond, Prunus dulcis (Miller) D. A. Webb, orchard in Kern Co., CA, during February and March 2002. Flight activity was measured by taking 1-min counts of bees exiting colonies on each of 9 d. Flight activity was best predicted with a model containing the effects of colony size (populations of adult bees and sealed brood), temperature, time of day, the interaction of adult bee population with temperature, and the interaction of adult bee population with time of day. Flight increased linearly with adult bee and brood population, had a quadratic relationship with temperature (increasing, but less so at higher temperatures), and had a quadratic relationship with time of day (decreasing, but less so at later times). Larger colonies had more response to changing temperatures and less response to different times of day than small colonies. Bee type had no direct influence on flight activity at any given colony size, temperature, or time of observation or when evaluated using a reduced data set retaining 34 Italian colonies and 32 Russian colonies whose mean sizes were equal. Overall, however, Russian colonies were less populous by about one-fourth and so fielded on average 71% of the foragers that Italian colonies did. Pollen collection was measured by capturing returning foragers on 4 d. The percentages of foragers with pollen were not different for the bee types.     

The indigenous honey bees of Saudi Arabia (Hymenoptera, Apidae, Apis mellifera jemenitica Ruttner): Their natural history and role in beekeeping

Apis mellifera jemenitica Ruttner (= yemenitica auctorum: videEngel 1999) has been used in apiculture throughout the Arabian Peninsula since at least 2000 BC. Existing literature demonstrates that these populations are well adapted for the harsh extremes of the region. Populations of Apis mellifera jemenitica native to Saudi Arabia are far more heat tolerant than the standard races often imported from Europe. Central Saudi Arabia has the highest summer temperatures for the Arabian Peninsula, and it is in this region where only Apis mellifera jemenitica survives, while other subspecies fail to persist. The indigenous race of Saudi Arabia differs from other subspecies in the region in some morphological, biological, and behavioral characteristics. Further taxonomic investigation, as well as molecular studies, is needed in order to confirm whether the Saudi indigenous bee populations represent a race distinct from Apis mellifera jemenitica, or merely an ecotype of this subspecies.     

A meta-analysis of effects of Bt crops on honey bees (Hymenoptera: Apidae)

Background

Honey bees (Apis mellifera L.) are the most important pollinators of many agricultural crops worldwide and are a key test species used in the tiered safety assessment of genetically engineered insect-resistant crops. There is concern that widespread planting of these transgenic crops could harm honey bee populations.

Methodology/Principal Findings

We conducted a meta-analysis of 25 studies that independently assessed potential effects of Bt Cry proteins on honey bee survival (or mortality). Our results show that Bt Cry proteins used in genetically modified crops commercialized for control of lepidopteran and coleopteran pests do not negatively affect the survival of either honey bee larvae or adults in laboratory settings.

Conclusions/Significance

Although the additional stresses that honey bees face in the field could, in principle, modify their susceptibility to Cry proteins or lead to indirect effects, our findings support safety assessments that have not detected any direct negative effects of Bt crops for this vital insect pollinator.     

Expression of Varroa sensitive hygiene (VSH) in commercial VSH honey bees (Hymenoptera: Apidae)

We tested six commercial sources of honey bees, Apis mellifera L. (Hymenoptera: Apidae), whose breeding incorporated the trait of Varroa sensitive hygiene (VSH). VSH confers resistance to the parasitic mite Varroa destructor Anderson & Trueman by enhancing the ability of the bees to hygienically remove mite-infested brood. VSH production queens (i.e., queens commercially available for use in beekeepers' production colonies) from the six sources were established in colonies which later were measured for VSH. Their responses were compared with those of colonies with three other types of queens, as follows: VSH queens from the selected closed population maintained by USDA-ARS for research and as a source of breeding germplasm, queens from the cooperating commercial distributor of this germplasm, and queens of a commercial, mite-susceptible source. The reduction of mite infestation in brood combs exposed to test colonies for 1 wk differed significantly between groups. On average, colonies with VSH production queens reduced infestation by 44%. This group average was intermediate between the greater removal by pure ARS VSH (76%) and the cooperators' breeding colonies (64%), and the lesser removal by susceptible colonies (7%). VSH production colonies from the different sources had variable expression of hygiene against mites, with average reduced infestations ranging from 22 to 74%. In addition, infertility was high among mites that remained in infested cells in VSH breeder colonies from ARS and the commercial distributor but was lower and more variable in VSH production colonies and susceptible colonies. Commercial VSH production colonies supply mite resistance that generally seems to be useful for beekeeping. Resistance probably could be improved if more VSH drones sources were supplied when VSH production queens are being mated.     

Host-seeking behaviour of tracheal mites (Acari: Tarsonemidae) on honey bees (Hymenoptera: Apidae)

The behaviour of the endoparasitic tracheal mite, Acarapis woodi (Rennie) on honey bees (Apis mellifera L.) is a challenge to observe because of its small size. Through a microscope, we videotaped this mite's movement on young bees, dead bees and bees exposed to vegetable oil. Previous studies have shown that solid vegetable oil decreases mite infestations in a bee colony. We hypothesized that the oil alters mite behaviour to the detriment of the parasite, thus helping to safeguard the host. Habitat-seeking behaviour, identified as necessary for mites to locate a new host environment, was disrupted on both dead and oil-treated bees. Questing behaviour, which is associated with transfer between hosts, increased significantly on the dead and oily bees. The behaviours of mites were significantly different between all three treatments (x ²=494.96, p<0.001 on dead bees and x ²=851.11, p<0.001 on oily bees). Both questing and seeking behaviours were significantly different on each of the thoracic treatments (F _2,66=7.88, p<0.001 and F _2,66=21.28, p<0.001) and mite questing behaviour was not altered between males and females on live or oily bees (F _1,22=0.25, p<0.62), but habitat seeking was (F _1,22=7.42, p<0.012). The male questing and habitat-seeking behaviours were observed. We conclude that oil-treated bees gained protection from habitat-seeking mites because the normal behaviour of the mites seeking an oviposition site is interrupted.     

An experiment on comb orientation by honey bees (Hymenoptera: Apidae) in traditional hives

The orientation of combs in traditional beehives is extremely important for obtaining a marketable honey product. However, the factors that could determine comb orientation in traditional hives and the possibilities of inducing honey bees, Apis mellifera (L.), to construct more desirable combs have not been investigated. The goal of this experiment was to determine whether guide marks in traditional hives can induce bees to build combs of a desired orientation. Thirty-two traditional hives of uniform dimensions were used in the experiment. In 24 hives, ridges were formed on the inner surfaces of the hives with fermented mud to obtain different orientations, circular, horizontal, and spiral, with eight replicates of each treatment. In the remaining eight control hives, the inner surface was left smooth. Thirty-two well-established honey bee colonies from other traditional hives were transferred to the prepared hives. The colonies were randomly assigned to the four treatment groups. The manner of comb construction in the donor and experimental hives was recorded. The results showed that 22 (91.66%) of the 24 colonies in the treated groups built combs along the ridges provided, whereas only 2 (8.33%) did not. Comb orientation was strongly associated with the type of guide marks provided. Moreover, of the 18 colonies that randomly fell to patterns different from those of their previous nests, 17 (94.4%) followed the guide marks provided, irrespective of the comb orientation type in their previous nest. Thus, comb orientation appears to be governed by the inner surface pattern of the nest cavity. The results suggest that even in fixed-comb hives, honey bees can be guided to build combs with orientations suitable to honey harvesting, without affecting the colonies.     

Miocene honey bees from the Randeck Maar of southwestern Germany (Hymenoptera, Apidae)

The Miocene Randeck Maar (southwestern Germany) is one of the only sites with abundant material of fossil honey bees. The fauna has been the focus of much scrutiny by early authors who recognized multiple species or subspecies within the fauna. The history of work on the Randeck Maar is briefly reviewed and these fossils placed into context with other Tertiary and living species of the genus Apis Linnaeus (Apinae: Apini). Previously unrecorded specimens from Randeck Maar were compared with earlier series in an attempt to evaluate the observed variation. A morphometric analysis of forewing venation angles across representative Recent and Tertiary species of Apis as well as various non-Apini controls was undertaken to evaluate the distribution of variation in fossil honey bees. The resulting dendrogram shows considerable variation concerning the wing venation of Miocene Apini, but intergradation of other morphological characters reveals no clear pattern of separate species. This suggests that a single, highly variable species was present in Europe during the Miocene. The pattern also supports the notion that the multiple species and subspecies proposed by earlier authors for the Randeck Maar honey bee fauna are not valid, and all are accordingly recognized as Apis armbrusteri Zeuner.     

Allozyme variation in bumble bees (Hymenoptera: Apidae)

Allozyme variation at an average of 37.3 loci was assessed in queens of 16 Bombus and 2 Psithyrus bumble bee species from North America. The mean expected heterozygosity (H) for the Bombus species was 0.008 +/- 0.006 (95% confidence limits) and that for the Psithyrus was 0.007 +/- 0.007. These levels are significantly lower than found in other Hymenoptera but are comparable to those found in previous studies of bumble bees based on far fewer loci. Neutral mutation and random genetic drift can account for the observed variation, but this implies a very small effective population size for species of bumble bees.