Suppression of queen rearing in European and Africanized honey beesApis mellifera L. by synthetic queen mandibular gland pheromone

Summary Queen rearing is suppressed in honey bees (Apis mellifera L.) by pheromones, particularly the queen's mandibular gland pheromone. In this study we compared this pheromonally-based inhibition between temperate and tropically-evolved honey bees. Colonies of European and Africanized bees were exposed to synthetic queen mandibular gland pheromone (QMP) for ten days following removal of resident queens, and their queen rearing responses were examined. Queen rearing was suppressed similarly in both European and Africanized honey bees with the addition of synthetic QMP, indicating that QMP acts on workers of both races in a comparable fashion. QMP completely suppressed queen cell production for two days, but by day six, cells containing queen larvae were present in all treated colonies, indicating that other signals play a role in the suppression of queen rearing. In queenless control colonies not treated with QMP, Africanized bees reared 30% fewer queens than Europeans, possibly due to racial differences in response to feedback from developing queens and/or their cells. Queen development rate was faster in Africanized colonies, or they selected older larvae to initiate cells, as only 1 % of queen cells were unsealed after 10 days compared with 12% unsealed cells in European colonies.     

Nest Defense Behavior in Colonies from Crosses Between Africanized and European Honey Bees (Apis mellifera L.) (Hymenoptera: Apidae)

Honey bee (Apis mellifera L.) colonies with either European or Africanized queens mated to European or Africanized drones alone or in combination were tested for defensive behavior using a breath test. The most defensive colonies were those with European or Africanized queens mated to Africanized drones. In colonies where both European and Africanized patrilines existed, most of the workers participating in nest defense behavior for the first 30 s after a disturbance were of African patrilines. Nest defense behavior appears to be genetically dominant in honey bees.     

Paternal effects on the defensive behavior of honeybees

The defensive behavior of 52 hybrid honeybee (Apis mellifera L.) colonies from four sets of crosses was studied and compared with that of European and Africanized bee colonies. Colonies containing F(1) hybrid workers were obtained through reciprocal crosses between European and Africanized bees. The total number of stings deposited by workers in a moving leather patch in 1 min was recorded. In each of the four sets of crosses, bees from hybrid colonies of Africanized paternity left more stings in leather patches than bees from hybrid colonies of European paternity. Results strongly suggest paternal effects of African origin increasing the defensive behavior of hybrid colonies. Although some degree of dominance was observed for high-defensive behavior in one of the four sets of crosses involving European paternity, most of the dominance effects reported in the literature appear to be the result of paternal effects. Several hypotheses to explain this phenomenon, as well as the implications of these effects on the fitness and breeding of honeybees are discussed.     

The effects of beta acids from hops (Humulus lupulus) on mortality of Varroa destructor (Acari: Varroidae)

Hop (Humulus lupulus L.) beta acids (HBA) were tested for miticidal effects on varroa destructor Anderson and Trueman, a parasitic mite of the honey bee (Apis mellifera L.). When varroa were placed on bees that had topical applications of 1?% HBA, there was 100?% mite mortality. Bee mortality was unaffected. Cardboard strips saturated with HBA and placed in colonies resulted in mite drop that was significantly greater than in untreated hives. HBA was detected on about 60?% of the bees in colonies during the first 48?h after application. Mite drop in colonies lasted for about 7?days with the highest drop occurring in the first 2–3?days after treatment. There was a reduction in the percentages of bees with HBA and in the amounts on their bodies after 7?days. Bee and queen mortality in the colonies were not affected by HBA treatments. When cardboard strips saturated with HBA were put in packages of bees, more than 90?% of the mites were killed without an increase in bee mortality. HBA might have potential to control varroa when establishing colonies from packages or during broodless periods.     

The influence of brood comb cell size on the reproductive behavior of the ectoparasitic mite Varroa destructor in Africanized honey bee colonies

Africanized honey bees (Apis mellifera, Hymenoptera: Apidae) in Brazil are tolerant of infestations with the exotic ectoparasitic mite, Varroa destructor (Mesostigmata: Varroidae), while the European honey bees used in apiculture throughout most of the world are severely affected. Africanized honey bees are normally kept in hives with both naturally built small width brood cells and with brood cells made from European-sized foundation, yet we know that comb cell size has an effect on varroa reproductive behavior. Three types (sizes) of brood combs were placed in each of six Africanized honey bee colonies: new (self-built) Africanized comb, new Italian comb (that the bees made from Italian-sized commercial foundation), and new Carniolan comb (built naturally by Carniolan bees). About 100 cells of each type were analyzed in each colony. The Africanized comb cells were significantly smaller in (inner) width (4.84 mm) than the European-sized comb cells (5.16 and 5.27 mm for Italian and Carniolan cells, respectively). The brood cell infestation rates (percentage cells infested) were significantly higher in the Carniolan-sized comb cells (19.3%) than in the Italian and Africanized cells (13.9 and 10.3%, respectively). The Carniolan-sized cells also had a significantly larger number of invading adult female mites per 100 brood cells (24.4) than did the Italian-sized cells (17.7) and the natural-sized Africanized worker brood cells (15.6). European-sized worker brood cells were always more infested than the Africanized worker brood cells in the same colony. There was a highly significant correlation (P<0.01) between cell width and the rate of infestation with varroa in four of the six colonies. The small width comb cells produced by Africanized honey bees may have a role in the ability of these bees to tolerate infestations by Varroa destructor, furthermore it appears that natural-sized comb cells are superior to over-sized comb cells for disease resistance.     

Population growth of Varroa destructor (Acari: Varroidae) in commercial honey bee colonies treated with beta plant acids

Varroa (Varroa destuctor Anderson and Trueman) populations in honey bee (Apis mellifera L.) colonies might be kept at low levels by well-timed miticide applications. HopGuard^® (HG) that contains beta plant acids as the active ingredient was used to reduce mite populations. Schedules for applications of the miticide that could maintain low mite levels were tested in hives started from either package bees or splits of larger colonies. The schedules were developed based on defined parameters for efficacy of the miticide and predictions of varroa population growth generated from a mathematical model of honey bee colony–varroa population dynamics. Colonies started from package bees and treated with HG in the package only or with subsequent HG treatments in the summer had 1.2–2.1 mites per 100 bees in August. Untreated controls averaged significantly more mites than treated colonies (3.3 mites per 100 bees). By October, mite populations ranged from 6.3 to 15.0 mites per 100 bees with the lowest mite numbers in colonies treated with HG in August. HG applications in colonies started from splits in April reduced mite populations to 0.12 mites per 100 bees. In September, the treated colonies had significantly fewer mites than the untreated controls. Subsequent HG applications in September that lasted for 3 weeks reduced mite populations to levels in November that were significantly lower than in colonies that were untreated or had an HG treatment that lasted for 1 week. The model accurately predicted colony population growth and varroa levels until the fall when varroa populations measured in colonies established from package bees or splits were much greater than predicted. Possible explanations for the differences between actual and predicted mite populations are discussed.     

Genetic dominance and worker interactions affect honeybee colony defense

Colonies of honeybees (Apis mellifera L.) were established thatvaried in the proportions of their workers that were of Europeanand hybrid (Africanized x European) descent. Colony defensiveresponses increased with higher proportions of hybrid workers.Colonies consisting exclusively of hybrid workers did not differin their response from "pure" Africanized colonies, suggestingthat the strong defensive behavior of Africanized workers isgenetically dominant. European workers became more defensivein colonies that also contained hybrid workers, whereas hybridworkers became less defensive in the same mixed colonies. Inmixed colonies hybrid workers were individually more likelythan Europeans to sting a leather target but not more likelyto guard the entrance.     

Is the number of antennal plate organs (sensilla placodea) greater in hygienic than in non-hygienic Africanized honey bees?

Hygienic behavior is a desirable trait in honey bees (Apis mellifera L.), as hygienic bees quickly remove diseased brood, interrupting the infectious cycle. Hygienic lines of honey bees appear to be more sensitive to the odors of dead and diseased honey bee brood, and Africanized honey bees are generally more hygienic than are European honey bees. We compared the number of sensilla placodea, antennal sensory structures involved in the perception of odor, in 10 bees from each of six hygienic and four non-hygienic colonies of Africanized honey bees. The sensilla placodea of three of the terminal segments (flagellomeres) of the right antenna of each bee were counted with a scanning electron microscope. There were no significant differences in the mean numbers of sensilla placodea between the hygienic and non-hygienic bees, though the variance was higher in the hygienic group. Flagellomere 4 had significantly more sensilla placodea than flagellomeres 6 and 8. However, there was no significant difference between the other two flagellomeres. As hygienic bees are capable of identifying dead, injured, or infested brood inside a capped brood cell, sensilla placodea probably have an important role in enabling worker bees to sense sick brood. However, we did not find greater numbers of this sensory structure in the antennae of hygienic, compared to non-hygienic Africanized honey bees.     

Comparative Sucrose Responsiveness in Apis mellifera and A. cerana Foragers

In the European honey bee, Apis mellifera, pollen foragers have a higher sucrose responsiveness than nectar foragers when tested using a proboscis extension response (PER) assay. In addition, Africanized honey bees have a higher sucrose responsiveness than European honey bees. Based on the biology of the Eastern honey bee, A. cerana, we hypothesized that A. cerana should also have a higher responsiveness to sucrose than A. mellifera. To test this hypothesis, we compared the sucrose thresholds of pollen foragers and nectar foragers in both A. cerana and A. mellifera in Fujian Province, China. Pollen foragers were more responsive to sucrose than nectar foragers in both species, consistent with previous studies. However, contrary to our hypothesis, A. mellifera was more responsive than A. cerana. We also demonstrated that this higher sucrose responsiveness in A. mellifera was not due to differences in the colony environment by co-fostering two species of bees in the same mixed-species colonies. Because A. mellifera foragers were more responsive to sucrose, we predicted that their nectar foragers should bring in less concentrated nectar compared to that of A. cerana. However, we found no differences between the two species. We conclude that A. cerana shows a different pattern in sucrose responsiveness from that of Africanized bees. There may be other mechanisms that enable A. cerana to perform well in areas with sparse nectar resources.     

The influence of nest site selection on the population dynamics of Africanized honey bees in an urban landscape

Urban landscapes provide habitat for many species, including domesticated and feral honey bees, Apis mellifera L. (Hymenoptera: Apidae). With recent losses of managed honey bee colonies, there is increasing interest in feral honey bee colonies and their potential contribution to pollination services in agricultural, natural, and urban settings. However, in some regions the feral honey bee population consists primarily of Africanized honey bees. Africanized honey bees (AHB) are hybrids between European honey bees and the African honey bee, Apis mellifera scutellataLepeletier, and have generated economic, ecological, and human health concerns because of their aggressive behavior. In this study, we used two long‐term datasets (7–10 years) detailing the spatial and temporal distribution of AHB colonies in Tucson, AZ, USA, where feral colonies occupy a variety of cavities including water meter boxes. A stage‐structured matrix model was used to elucidate the implications of nest site selection and the effects of colony terminations on the structure and dynamics of the AHB population. Our results suggest that Tucson's AHB population is driven by a relatively small number of ‘source’ colonies that escape termination (ca. 0.165 colonies per km² or 125 colonies in total), although immigrating swarms and absconding colonies from the surrounding area may have also contributed to the stability of the Tucson AHB population. Furthermore, the structure of the population has likely been impacted by the number and spatial distribution of water meter boxes across the city. The study provides an example of how urban wildlife populations are driven by interactions among landscape structure, human management, and behavioral traits conferred by an invasive genotype.     

Gram-positive cocci from apiarian sources

Frass from the greater wax moth, Galleria mellonella, obtained from feral colonies of honey bees, Apis mellifera; from domesticated (managed) honey bee colonies; and from a laboratory culture of the wax moth was sampled for Gram-positive cocci. One hundred twenty-three of these organisms were isolated and identified. Frass from domesticated colonies yielded only one isolate. Equal numbers of isolates (61) were obtained from frass from feral bee colonies and from the wax moth culture. Catalase-negative cocci were predominant in frass from feral colonies, whereas catalase-positive cocci were the most common isolates from frass from the wax moth culture. Catalase-positive cocci were identified as Staphylococcus epidermidis and Micrococcus sp. Catalase-negative cocci were Streptococcus faecalis var. faecalis and S. faecium. These results are discussed in relation to the rarity of Gram-positive cocci associated with honey bees, pollen, and nectar in Arizona and the frequency of association with honey bees and wax moth frass of bacteria resembling Arthrobacter spp. that appear as Gram-positive cocci during one stage of the life cycle.     

How Honey Bee Colonies Survive in the Wild: Testing the Importance of Small Nests and Frequent Swarming

The ectoparasitic mite, Varroa destructor, and the viruses that it transmits, kill the colonies of European honey bees (Apis mellifera) kept by beekeepers unless the bees are treated with miticides. Nevertheless, there exist populations of wild colonies of European honey bees that are persisting without being treated with miticides. We hypothesized that the persistence of these wild colonies is due in part to their habits of nesting in small cavities and swarming frequently. We tested this hypothesis by establishing two groups of colonies living either in small hives (42 L) without swarm-control treatments or in large hives (up to 168 L) with swarm-control treatments. We followed the colonies for two years and compared the two groups with respect to swarming frequency, Varroa infesttion rate, disease incidence, and colony survival. Colonies in small hives swarmed more often, had lower Varroa infestation rates, had less disease, and had higher survival compared to colonies in large hives. These results indicate that the smaller nest cavities and more frequent swarming of wild colonies contribute to their persistence without mite treatments.     

POPULATION DIFFERENTIATION AND RACIAL ADMIXTURE IN THE AFRICANIZED HONEYBEE (APIS MELLIFERA L.)

To study the degree of interpopulational differentiation and racial admixture in Africanized honeybees, we collected worker bees from three regions of Brazil (the northeast, the state of Sao Paulo, and Porto Alegre) and from Uruguay and determined their genotypes for 10 enzyme loci. We also performed a morphometric analysis on forewing measurements of worker bees from the northeast and Porto Alegre regions of Brazil and from Paysandu, Uruguay. Comparative analysis of interpopulational heterogeneity snowed that there are significant differences, especially at the Mdh locus, among the populations from different regions. An increase in the frequency of the Mdh^B allele was observed from north to south, with predominance in the Uruguayan populations. A small component of interpopulational variability was detected in the populations studied. Racial admixture was calculated from information obtained for Mdh in Africa and Europe. The percentages of racial admixture differed slightly but significantly among Brazilian regions. The morphometric study based on canonical variables exhibited a similar pattern. The greater proportions of Apis mellifera adansonii alleles in the admixture may be explained by selection during the initial stage of migration of Africanized bees and by preferential mating between individuals of the same race. Differences in the proportions of A. m. adansonii alleles between regions indicate incipient populational differentiation of Africanized bees. We suggest that greater gene flow from the European races in the south of Brazil could be one of the causes of this phenomenon.     

A push‐pull integrated pest management scheme for preventing use of parrot nest boxes by invasive Africanized honey bees

Africanized honey bees (Apis mellifera scutellata) compete with endangered parrots for nest boxes and can hamper conservation efforts. We tested an integrated pest management push‐pull protocol in the Atlantic Forest in São Paulo, Brazil, in an effort to prevent bee swarms from colonizing nest boxes (N = 30 in the forest plus five in aviaries) meant for use by Vinaceous‐breasted Amazons (Amazona vinacea). Fifteen parrot nest boxes were treated with a permethrin insecticide to “push” scout bees away and each parrot box was paired with a bee trap box containing a pheromone lure to “pull” bees. Over a 1‐yr period (March 2013 to March 2014), 29 insect colonies moved into 18 of the 35 trap boxes. Nine Africanized honey bee, three native Jatai bee (Tetragonisca sp.), and 17 wasp colonies occupied trap boxes. Only one experimental push‐pull pair untreated parrot box was invaded by bees and no parrot boxes in aviaries were colonized. Four of the parrot nest boxes were occupied by birds during our study. Although none were used by Vinaceous‐breasted Amazons, Southern House Wrens (Troglodytes musculus), Green‐winged Saltators (Saltator similis), and Plain Parakeets (Brotogeris tirica) nested in the boxes and all nests were successful. Although long‐term studies are needed before drawing conclusions about the effectiveness of trap boxes, our results suggest that a push‐pull protocol may prove useful for reducing the use of nest boxes meant for parrots and other cavity‐nesting birds by Africanized honey bees and other insects.     

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.     

Genetic variation in worker temporal polyethism and colony defensiveness in the honey bee, Apis mellifera

To test the hypothesis that colonies of honey bees composedof workers with faster rates of adult behavioral developmentare more defensive than colonies composed of workers with slowerbehavioral development, we determined whether there is a correlationbetween genetic variation in worker temporal polyethism andcolony defensiveness. There was a positive correlation for thesetwo traits, both for European and Africanized honey bees. Thecorrelation was larger for Africanized bees, due to differencesbetween Africanized and European bees, differences in experimentaldesign, or both. Consistent with these results was the findingthat colonies with a higher proportion of older bees were moredefensive than colonies of the same size that had a lower proportionof older bees. There also was a positive correlation betweenrate of individual behavioral development and the intensityof colony flight activity, and a negative correlation betweencolony defensiveness and flight activity. This suggests thatthe relationship between temporal polyethism and colony defensivenessmay vary with the manner in which foraging and defense dutiesare allocated among a colony's older workers. These resultsindicate that genotypic differences in rates of worker behavioraldevelopment can influence the phenotype of a honey bee colonyin a variety of ways.     

Viral Infection Affects Sucrose Responsiveness and Homing Ability of Forager Honey Bees,Apis mellifera L.

Honey bee health is mainly affected by Varroa destructor, viruses, Nosema spp., pesticide residues and poor nutrition. Interactions between these proposed factors may be responsible for the colony losses reported worldwide in recent years. In the present study, the effects of a honey bee virus, Israeli acute paralysis virus (IAPV), on the foraging behaviors and homing ability of European honey bees (Apis mellifera L.) were investigated based on proboscis extension response (PER) assays and radio frequency identification (RFID) systems. The pollen forager honey bees originated from colonies that had no detectable level of honey bee viruses and were manually inoculated with IAPV to induce the viral infection. The results showed that IAPV-inoculated honey bees were more responsive to low sucrose solutions compared to that of non-infected foragers. After two days of infection, around 10⁷ copies of IAPV were detected in the heads of these honey bees. The homing ability of IAPV-infected foragers was depressed significantly in comparison to the homing ability of uninfected foragers. The data provided evidence that IAPV infection in the heads may enable the virus to disorder foraging roles of honey bees and to interfere with brain functions that are responsible for learning, navigation, and orientation in the honey bees, thus, making honey bees have a lower response threshold to sucrose and lose their way back to the hive.     

Genotype-environment interactions in honeybee guarding behaviour

Honeybees have an age-based division of labour that is influenced by genetic variability for the tendency to perform specific tasks. Individuals in a honeybee colony comprise diverse genotypes and their interactions can influence task allocation. Colonies from an African race (Africanized honeybees, AHB, Apis mellifera scutellata Ruttner) usually produce a much stronger defensive response than do European races of honeybees (EHB), and these races may differ in how individuals are allocated to the tasks of guarding and stinging. We observed guarding behaviour in colony environments that varied in proportions of genotypes (AHB, EHB) and population size. In large colonies, AHB showed much greater guarding persistence (number of days guarding) than EHB; hybrids were intermediate. In another series of experiments, three families each of AHB and EHB were cofostered in colonies with different AHB: EHB ratios, then tested in large and small colonies. In colonies of both sizes, colony environment interacted with both famly and type (AHB or EHB) for propensity to guard. Individuals of both types guarded more persistently in large colonies, but family and type both interacted with environment. EHB were more likely to initiate guarding bouts in low-AHB colonies, but persistence did not change with environment. AHB were insensitive to effects of environment for the tendency to initiate guarding behaviour, but were more persistent in high-AHB environments. EHB and AHB may differ in how they allocate individuals to guarding. The positive reinforcement of behaviour that occurs in high-defensive environments and in large populations could cause a stronger stinging response through alarm pheromone recruitment. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Uncapping activity of Apis mellifera L. (Hymenoptera: Apidae) towards worker brood cells infested with the mite Varroa destructor Anderson & Treuman (Mesostigmata: Varroidae)

Varroosis, a disease caused by the mite Varroa destructor Anderson and Treuman has killed hundreds of thousands of Apis mellifera L. colonies in various parts of the world. Nevertheless, the damage caused by this mite varies with the type of bee and climate conditions. Varroa causes little damage to Africanized bee colonies in Brazil, as the infestation rates are relatively stable and low. We evaluated the hygienic behavior (uncapping and removal of brood) of highly hygienic Africanized bees using combs with worker brood cells infested (naturally) and no infested with V. destructor. The daily uncapping rate, measured in eight colonies during six days, was 3.5 fold higher in the combs infested with varroa compared to no infested combs. The results show that the Africanized bees are able to recognise and remove brood cells naturally infested with V. destructor what is an important mechanism for tolerance against varroa.     

Comparison of the activity and productivity of Carniolan (Apis mellifera carnica Pollmann) and Yemeni (Apis mellifera jemenitica Ruttner) subspecies under environmental conditions of the Al-Ahsa oasis of eastern Saudi Arabia

This study was conducted at the apiary of the Agricultural and Veterinary Training and Research Station of King Faisal University in the Al-Ahsa oasis of eastern Saudi Arabia. We performed a comparison between Carniolan (Apis mellifera carnica Pollmann) and Yemeni (Apis mellifera jemenitica Ruttner) honeybee races to determine the monthly fluctuations in foraging activity, pollen collection, colony growth and honey yield production under the environmental conditions of the Al-Ahsa oasis of eastern Saudi Arabia. We found three peaks in the flight activity of the two races, and the largest peaks occurred during September and October. Compared to Carniolan bee colonies, the performance of Yemeni bee colonies was superior in terms of stored pollen, worker and drone brood rearing, and the adult population size. The Carniolan bee colonies produced 27.77% and 27.50% more honey than the Yemeni bee colonies during the flow seasons of alfalfa and sidir, respectively, with an average increase of 27.64%. It could be concluded that the race of bees is an important factor affecting the activity and productivity of honeybee colonies. The Yemeni bee race produced more pollen, a larger brood and more bees, which exhibited a longer survival. The imported Carniolan bees can be reared in eastern Saudi Arabia, but the Yemeni bee race is still better.