Social parasitism by Cape honeybee workers in colonies of their own subspecies (Apis mellifera capensis Esch.)

Social parasitism is widespread in the eusocial insects. Although social parasites often show a reduced worker caste, unmated workers can also parasitize colonies. Cape honeybee workers, Apis mellifera capensis, can establish themselves as social parasites in host colonies of other honeybee subspecies. However, it is unknown whether social parasitism by laying workers also occurs among Cape honeybee colonies. In order to address this question we genotyped worker offspring of six queenless A. m. capensis colonies and determined the maternity of the reproducing workers. We found that three non-nestmate workers dominated reproduction in a host colony and produced 62.5% of the progeny. Our results show that social parasitism by laying workers is a naturally occurring part of the biology of Cape honeybees. However, such social parasitism is not frequently found (6.41% of the total worker offspring) probably due to co-evolutionary processes among A. m. capensis resulting in an equilibrium between selection for reproductive dominance in workers, colony maintenance and queen adaptation. Received 28 July 2005; revised 19 September and 11 November 2005; accepted 16 November 2005.     

Cape honeybees, Apis mellifera capensis, police worker-laid eggs despite the absence of relatedness benefits

In the Cape honeybee, Apis mellifera capensis, workers lay diploid(female) eggs via thelytoky. In other A. mellifera subspecies,workers lay haploid (male) eggs via arrhenotoky. When thelytokousworker reproduction occurs, worker policing has no relatednessbenefit because workers are equally related to their sisterworkers' clonal offspring and their mother queen's female offspring.We studied worker policing in A. m. capensis and in the arrhenotokousAfrican honeybee A. m. scutellata by quantifying the removalrates of worker-laid and queen-laid eggs. Discriminator coloniesof both subspecies policed worker-laid eggs of both their ownand the other subspecies. The occurrence of worker policing,despite the lack of relatedness benefit, in A. m. capensis stronglysuggests that worker reproduction is costly to the colony andthat policing is maintained because it enhances colony efficiency.In addition, because both subspecies policed each others eggs,it is probable that the mechanism used in thelytokous A. m.capensis to discriminate between queen-laid and worker-laideggs is the same as in arrhenotokous A. m. scutellata.     

Male fitness of honeybee colonies (Apis mellifera L.)

Honeybees (Apis mellifera L.) have an extreme polyandrous mating system. Worker offspring of 19 naturally mated queens was genotyped with DNA microsatellites, to estimate male reproductive success of 16 drone producing colonies. This allowed for estimating the male mating success on both the colony level and the level of individual drones. The experiment was conducted in a closed population on an isolated island to exclude interferences of drones from unknown colonies. Although all colonies had produced similar numbers of drones, differences among the colonies in male mating success exceeded one order of magnitude. These differences were enhanced by the siring success of individual drones within the offspring of mated queens. The siring success of individual drones was correlated with the mating frequency at the colony level. Thus more successful colonies not only produced drones with a higher chance of mating, but also with a significantly higher proportion of offspring sired than drones from less successful colonies. Although the life cycle of honeybee colonies is very female centred, the male reproductive success appears to be a major driver of natural selection in honeybees.     

Preparation for disturbance-induced absconding of Cape honeybee colonies (Apis mellifera capensis Esch.)

African honeybees, Apis mellifera, are characterised by frequent disturbance-induced absconding. However, the effectiveness in preparation before such disturbance-induced absconding has not been rigorously quantified yet. We investigated the effectiveness of preparation for disturbance-induced absconding by evaluating colony phenotypes prior to and after absconding in ten colonies of the Cape honeybee, A. m. capensis. Seven non-absconding colonies at the same apiary were used as controls. While seven absconded colonies left neither stores nor brood behind, three colonies abandoned only a small area of honey, pollen, open or capped brood. At the end of the observations, the control colonies still had pollen and honey stores and brood. The mean reduction rate between a major disturbance and the absconding event was 0.052 ± 0.018 cm² stores and open brood per worker per day. Our results demonstrate that disturbance-induced absconding can also occur with preparation, if the disturbance is not highly destructive and enough time for preparation is available. We conclude that Cape honeybee colonies can show a considerable high effectiveness in their preparation before disturbance-induced absconding, which limits the loss of colony resources. In light of the general high mobility of African honeybee colonies such an efficient behaviour is probably adaptive. Received 22 December 2004; revised 3 June 2005; accepted 13 June 2005.     

Anarchy in the UK: Detailed genetic analysis of worker reproduction in a naturally occurring British anarchistic honeybee,Apis mellifera,colony using DNA microsatellites

Anarchistic behaviour is a very rare phenotype of honeybee colonies. In an anarchistic colony, many workers' sons are reared in the presence of the queen. Anarchy has previously been described in only two Australian colonies. Here we report on a first detailed genetic analysis of a British anarchistic colony. Male pupae were present in great abundance above the queen excluder, which was clearly indicative of extensive worker reproduction and is the hallmark of anarchy. Seventeen microsatellite loci were used to analyse these male pupae, allowing us to address whether all the males were indeed workers' sons, and how many worker patrilines and individual workers produced them. In the sample, 95 of 96 of the males were definitely workers' sons. Given that approximately 1% of workers' sons were genetically indistinguishable from queen's sons, this suggests that workers do not move any queen-laid eggs between the part of the colony where the queen is present to the area above the queen excluder which the queen cannot enter. The colony had 16 patrilines, with an effective number of patrilines of 9.85. The 75 males that could be assigned with certainty to a patriline came from 7 patrilines, with an effective number of 4.21. They were the offspring of at least 19 workers. This is in contrast to the two previously studied Australian naturally occurring anarchist colonies, in which most of the workers' sons were offspring of one patriline. The high number of patrilines producing males leads to a low mean relatedness between laying workers and males of the colony. We discuss the importance of studying such colonies in the understanding of worker policing and its evolution.     

Social parasitism by honeybee workers (Apis mellifera capensis Escholtz): host finding and resistance of hybrid host colonies

We studied possible host finding and resistance mechanisms ofhost colonies in the context of social parasitism by Cape honeybee(Apis mellifera capensis) workers. Workers often join neighboringcolonies by drifting, but long-range drifting (dispersal) tocolonies far away from the maternal nests also rarely occurs.We tested the impact of queenstate and taxon of mother andhost colonies on drifting and dispersing of workers and on the hosting of these workers in A. m. capensis, A. m. scutellata,and their natural hybrids. Workers were paint-marked accordingto colony and reintroduced into their queenright or queenlessmother colonies. After 10 days, 579 out of 12,034 labeled workerswere recaptured in foreign colonies. We found that driftingand dispersing represent different behaviors, which were differentlyaffected by taxon and queenstate of both mother and host colonies.Hybrid workers drifted more often than A. m. capensis and A.m. scutellata. However, A. m. capensis workers dispersed moreoften than A. m. scutellata and the hybrids combined, and A. m. scutellata workers also dispersed more frequently than thehybrids. Dispersers from queenright A. m. capensis colonieswere more often found in queenless host colonies and vice versa,indicating active host searching and/or a queenstate-discriminatingguarding mechanism. Our data show that A. m. capensis workersdisperse significantly more often than other races of A. mellifera,suggesting that dispersing represents a host finding mechanism.The lack of dispersal in hybrids and different hosting mechanismsof foreign workers by hybrid colonies may also be responsiblefor the stability of the natural hybrid zone between A. m.capensis and A. m. scutellata.     

Maintenance and loss of heterozygosity in a thelytokous lineage of honey bees (Apis mellifera capensis)

An asexual lineage that reproduces by automictic thelytokous parthenogenesis has a problem: rapid loss of heterozygosity resulting in effective inbreeding. Thus, the circumstances under which rare asexual lineages thrive provide insights into the trade-offs that shape the evolution of alternative reproductive strategies across taxa. A socially parasitic lineage of the Cape honey bee, Apis mellifera capensis, provides an example of a thelytokous lineage that has endured for over two decades. It has been proposed that cytological adaptations slow the loss of heterozygosity in this lineage. However, we show that heterozygosity at the complementary sex determining (csd) locus is maintained via selection against homozygous diploid males that arise from recombination. Further, because zygosity is correlated across the genome, it appears that selection against diploid males reduces loss of homozygosity at other loci. Selection against homozygotes at csd results in substantial genetic load, so that if a thelytokous lineage is to endure, unusual ecological circumstances must exist in which asexuality permits such a high degree of fecundity that the genetic load can be tolerated. Without these ecological circumstances, sex will triumph over asexuality. In A. m. capensis, these conditions are provided by the parasitic interaction with its conspecific host, Apis mellifera scutellata.     

Egg marking pheromones of anarchistic worker honeybees (Apis mellifera)

In honeybees, worker policing via egg eating enforces functionalworker sterility in colonies with a queen and brood. It is thoughtthat queens mark their eggs with a chemical signal, indicatingthat their eggs are queen-laid. Worker-laid eggs lack this signaland are, therefore, eaten by policing workers. Anarchistic workerhoneybees have been hypothesized to circumvent worker policingby mimicking the queen egg-marking signal. We investigated thisphenomenon by relating chemical profiles of workers and theireggs to egg acceptability. We found that the ability of someworkers (anarchistic workers in queenright colonies and deviantworkers from a queenless colony) to lay more acceptable eggsis due to them producing significant amounts of queen-like estersfrom their Dufour's gland. These esters appear to be transferredto eggs during laying and increase egg survival. However, theseesters cannot be the normal queen egg-marking signal, as theyare generally absent from queen-laid eggs and only increasethe short-term persistence of worker-laid eggs, because only7–30% of anarchistic worker-laid eggs persisted to hatchingversus 91–92% of queen-laid eggs. All workers can producesome esters, but only workers that greatly increase their esterproduction lay more acceptable eggs. The production of estersappears to be a flexible response, as anarchistic workers rearedin queenless colonies did not increase their ester production,while some deviant workers in queenless colonies did increasetheir ester production.     

What the honeybee sees: a review of the recognition system of Apis mellifera

Abstract. For many years, two opposing theories have dominated our ideas of what honeybees see. The earliest proposal based on training experiments was that bees detected only simple attributes or features, irrespective of the actual pattern. The features demonstrated experimentally before 1940 were the disruption of the pattern (related to spatial frequency), the area of black or colour, the length of edge, and the angle of orientation of a bar or grating. Cues discovered recently are the range, and radial and tangential edges, and symmetry, relative to the fixation point, which is usually the reward hole. This theory could not explain why recognition failed when the pattern was moved. In the second theory, proposed in 1969, the bee detected the retinotopic directions of black or coloured areas, and estimated the areas of overlap and nonoverlap on each test pattern with the corresponding positions in the training pattern. This proposal explained the progressive loss of recognition as a test pattern was moved or reduced in size, but required that the bees saw and remembered the layout of every learned pattern and calculated the mismatch with each test image. Even so, the same measure of the mismatch was given by many test patterns and could not detect a pattern uniquely. Moreover, this theory could not explain the abundant evidence of simple feature detectors. Recent work has shown that bees learn one or more of a limited number of simple cues. A newly discovered cue is the position, mainly in the vertical direction, of the common centre (centroid) of black areas combined together. Significantly, however, the trained bees look for the cues mentioned above only in the range of places where they had occurred during the training. These two observations made possible a synthesis of both theories. There is no experimental evidence that the bees detect or re-assemble the layout of patterns in space; instead, they look for a cue in the expected place. With an array of detectors of the known cues, together with their directions, this mechanism would enable bees to recognize each familiar place from the coincidences of cues in different directions around the head.     

意大利蜜蜂工蜂卵巢发育的研究

蜜蜂Apis mellifera L.蜂群中的工蜂卵巢发育和工蜂产卵现象受多种因素控制,了解其影响因素对养蜂生产具有重要意义。本研究将意大利蜜蜂蜂群设置为囚王群(tg1)、无王有子群(tg2)、无王无子群(tg3)以及正常有王蜂群(CK)4个试验组,通过对工蜂卵巢管的显微观察,确定不同处理组工蜂在不同时间段内卵巢的发育情况。结果表明:随着时间的延长,与CK处理组相比,tg3处理组中的工蜂卵巢发育水平最高,tg2次之,tg1最低;在31d时,4个处理组两两之间差异均达到显著水平(P<0.05)。tg1、tg2和tg3处理组中工蜂产卵前期时间分别为35、22和17d,而CK蜂群在试验期内未出现工蜂产卵现象;tg2和tg3处理组的工蜂产卵的封盖前期时间分别为8和6d,而tg1和CK组在试验期内未出现子房封盖现象。蜂群失王时间过长会刺激工蜂卵巢发育,并导致其产卵;蜂群的短期失王和蜂王老化也会刺激工蜂卵巢发育,但是刺激程度较低;蜂群中的蜂子能抑制工蜂卵巢管的发育,因此在蜂群短时间失王时可以适当地补充子脾延缓工蜂卵巢发育。     

The Releaser Effects of the Tergal Gland Secretion of Queen Honeybees (Apis mellifera)

Queen honeybees are attractive to their workers, due partially to the pheromonal bouquet they secrete. Queen mandibular gland pheromone is a powerful attractant to worker honeybees but it is not solely responsible for eliciting retinue behavior. The attractiveness of virgin queen tergal gland secretions and queen mandibular pheromone to worker honeybees was tested using a retinue bioassay. The number of workers attending the treated pseudoqueen lures was videorecorded in order to allow for the quantification of attractiveness. Queen mandibular gland secretions were more attractive than tergal gland secretions (P 0.008), and both queen tergal gland secretions (P 0.0002) and mandibular gland secretions (P 0.0001) were significantly more attractive than the control treatment. This laboratory bioassay indicates that queen tergal gland secretions have a releaser effect that evokes retinue behavior from worker honeybees.     

蜜蜂蜂王与工蜂级型分化研究进展

蜜蜂ApismekiferaL .是典型的社会性昆虫 ,蜂王和工蜂都是由受精卵发育而来的二倍体成蜂 ,但是在形态、生理、行为等方面有明显的差异 ,属于不同的级型。蜂王和工蜂的级型分化的关键时期发生在幼虫的 4龄末至 5龄止。分化是由分化基因调控的 ,幼虫期食物的质和量是分化的外部决定因子。JH对两级型中卵巢的分化有非常重要的调控作用。蜜蜂脑或其它组织中可能有分泌调控CA的咽侧体调节激素 ,它们通过对CA中JH的合成和分泌的调控而参与了分化的调控。章鱼胺等生物胺也参与了分化调控过程。     

Visual discrimination by the honeybee (Apis mellifera): the position of the common centre as the cue

Abstract. Bees can be trained to discriminate between a target with a 20° spot above a 10° spot of the same colour, and another target with the spots exchanged in position. Tests show that they do not remember the separate positions of spots of the same colour (including black) on the same target. The bees discriminate the difference in positions, in the vertical direction, of the common centres of the spots taken together, with or without green contrast.
Similar results are obtained in discriminations of a fixed T shape, each composed of two broad black bars subtending 8 by 24°, vs the same shape inverted. The trained bees fail to discriminate between the T shapes when the centroids are at the same level in the vertical direction. Moving the shapes in the horizontal direction in tests has less effect. Quite different results are obtained when the two bars of the T shape differ in colour. The bees discriminate the positions of the two colours separately, but they still fail to discriminate the shape of the T. The results can be explained by filters that detect the intensities within their fields, irrespective of shape, and weigh them according to their vertical angles from the horizontal midline. The normal function of these filters could be to detect the levels of objects relative to the horizon when the bee is in flight.     

狄斯瓦螨和蜜蜂病毒对意大利蜜蜂蜂群越冬的影响

越冬期是蜂群损失最主要的阶段.通过比较分析45个意大利蜜蜂Apis mellifera ligustica蜂群在繁殖越冬蜂前的狄斯瓦螨Varroa destructor寄生率和病毒感染情况、越冬表现及越冬期存活蜂群的病毒感染情况等,探究与越冬期蜜蜂健康紧密相关的影响因素.结果表明,繁殖越冬蜂前蜂群的狄斯瓦螨寄生率与蜜蜂...     

Natural selection of Varroa jacobsoni explains the different reproductive strategies in colonies of Apis cerana and Apis mellifera

In colonies of European Apis mellifera, Varroa jacobsoni reproduces both in drone and in worker cells. In colonies of its original Asian host, Apis cerana, the mites invade both drone and worker brood cells, but reproduce only in drone cells. Absence of reproduction in worker cells is probably crucial for the tolerance of A. cerana towards V. jacobsoni because it implies that the mite population can only grow during periods in which drones are reared. To test if non-reproduction of V. jacobsoni in worker brood cells of A. cerana is due to a trait of the mites or of the honey-bee species, mites from bees in A. mellifera colonies were artificially introduced into A. cerana worker brood cells and vice versa. Approximately 80% of the mites from A. mellifera colonies reproduced in naturally infested worker cells as well as when introduced into worker cells of A. mellifera and A. cerana. Conversely, only 10% of the mites from A. cerana colonies reproduced, both in naturally infested worker cells of A. cerana and when introduced into worker cells of A. mellifera. Hence, absence of reproduction in worker cells is due to a trait of the mites. Additional experiments showed that A. cerana bees removed 84% of the worker brood that was artificially infested with mites from A. mellifera colonies. Brood removal started 2 days after artificial infestation, which suggests that the bees responded to behaviour of the mites. Since removal behaviour of the bees will have a large impact on fitness of the mites, it probably plays an important role in selection for differential reproductive strategies. Our findings have large implications for selection programmes to breed less-susceptible bee strains. If differences in non-reproduction are mite specific, we should not only look for non-reproduction as such, but for colonies in which non-reproduction in worker cells is selected. Hence, in selection programmes fitness of mites that reproduce in both drone and worker cells should be compared to fitness of mites that reproduce only in drone cells. © Rapid Science Ltd. 1998     

Adaptation of worker honeybees (Apis mellifera) to their alarm pheromones

ABSTRACT. Honeybees rapidly became adapted to synthetic alarm pheromone components dispensed within their hives, and were less; inclined to sting both unmarked targets and targets marked with the; alarm pheromone. The possible application to beekeeping is discussed.     

意大利工蜂前胃显微结构观察

通过解剖镜和扫描电子显微镜对意大利工蜂Apismellifera ligusticaL.前胃的形态结构进行了观察,发现前胃包括前胃瓣和前胃管两部分。前胃瓣是由4个瓣片环围而成,球形,在蜜囊中,每个瓣片的长度约为0.8mm,前端为三角形结构,可以运动,边长约为0.35mm,三角形的前缘有较长的毛形结构。在每个三角形顶端各有一个直径约为50μm瘤形突起。前胃瓣最大孔径为0.89～1.00mm,具有调整和控制食物进入中肠的功能。前胃管是一直径0.01～0.02mm的较为柔软的管道,插入中肠5mm,该结构可防止中肠食物的倒流。前胃瓣上有7种毛状结构,分布在前胃瓣的不同部位,该结构的功能是分离和过滤、捕捉和感觉功能。本研究为理解工蜂前胃在消化系统中的功能和作用以及昆虫形态学和昆虫分类学提供理论依据。     

Bacterial diversity in worker adults of Apis mellifera capensis and Apis mellifera scutellata (Insecta: Hymenoptera) assessed using 16S rRNA sequences

High-fidelity PCR of 16S rRNA sequences was used to identify bacteria associated with worker adults of the honeybee subspecies Apis mellifera capensis and Apis mellifera scutellata. An expected approximately 1.5-kb DNA band, representing almost the entire length of the 16S rRNA gene, was amplified from both subspecies and cloned. Ten unique sequences were obtained: one sequence each clustered with Bifidobacterium (Gram-positive eubacteria), Lactobacillus (Gram-positive eubacteria), and Gluconacetobacter (Gram-negative alpha-proteobacteria); two sequences each clustered with Simonsiella (beta-proteobacteria) and Serratia (gamma-proteobacteria); and three sequences each clustered with Bartonella (alpha-proteobacteria). Although the sequences relating to these six bacterial genera initially were obtained from either A. m. capensis or A. m. scutellata or both, newly designed honeybee-specific 16S rRNA primers subsequently amplified all sequences from all individual workers of both subspecies. Attempts to amplify these sequences from eggs have failed. However, the wsp primers designed to amplify Wolbachia DNA from arthropods, including these bees, consistently produced a 0.6-kb DNA band from individual eggs, indicating that amplifiable bacterial DNA was present. Hence, the 10 bacteria could have been acquired orally from workers or from other substrates. This screening of 16S rRNA sequences from A. m. capensis and A. m. scutellata found sequences related to Lactobacillus and Bifidobacterium which previously had been identified from other honeybee subspecies, as well as sequences related to Bartonella, Gluconacetobacter, Simonsiella/Neisseria, and Serratia, which have not been identified previously from honeybees.     

Effect of triflumuron on brood development and colony survival of free-flying honeybee, Apis mellifera L.

Abstract:  The effect of the insect growth regulator (IGR) triflumuron (Alsystin^® 25 WP) on honeybee, Apis mellifera L. (Hym., Apidae), was studied in a semi-field test. Free-living colonies were fed one litre per hive of sucrose syrup containing 0, 0.025, 0.25 or 2.5 g of triflumuron. A significant reduction in flight activity was noted 6–10 weeks post-treatment at the two higher doses. These colonies reared less brood than before treatment. While the comb area occupied by uncapped brood was as high as [0.025 and 0.25 g active ingredient (a.i.)] or higher (2.5 g a.i.) than before treatment, there was a significant decline in capped brood at the two higher doses, indicating enhanced larval mortality. No capped brood was reared in the hive treated at the highest dose from 3–9 weeks post-treatment. Yet there was a significant accumulation of pollen and honey in the brood compartment at all doses. All colonies except the one treated at the highest dose survived the following winter. However, at 43 weeks post-treatment, hives treated at intermediate and low doses showed a significant increase in uncapped brood and a significant decrease in capped brood. This study revealed a strong residual toxicity of triflumuron to brood and substantiated its classification as hazardous to honeybee.     

Cheaters sometimes prosper: targeted worker reproduction in honeybee (Apis mellifera) colonies during swarming

Kin selection theory predicts that honeybee (Apis mellifera) workers should largely refrain from producing their own offspring, as the workers collectively have higher inclusive fitness if they rear the sons of their mother, the queen. Studies that have quantified levels of ovary activation and reproduction among workers have largely supported this prediction. We sampled pre‐emergent male pupae and adult workers from seven colonies at regular intervals throughout the reproductive part of the season. We show that the overall contribution of workers to male (drone) production is 4.2%, nearly 40 times higher than is generally reported, and is highest during reproductive swarming, when an average of 6.2% of the males genotyped are worker‐produced. Similarly, workers in our samples were 100 times more likely to have active ovaries than previously assumed. Worker reproduction is seasonally influenced and peaks when colonies are rearing new queens. Not all worker subfamilies contribute equally to reproduction. Instead, certain subfamilies are massively over‐represented in drone brood. By laying eggs within the period in which many colonies produce virgin queens, these rare worker subfamilies increase their direct fitness via their well‐timed sons.