Studies on mixed-species colonies of honeybees, Apis cerana and Apis mellifera
- Authors: Yang, Ming-Xian
- Date: 2010
- Subjects: Bees Apis cerana Honeybee Honeybee -- Behavior Bee culture Honeybee -- Physiology Insect societies Animal communication Bees -- Nests
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5779 , http://hdl.handle.net/10962/d1005467
- Description: The honeybees Apis cerana and Apis mellifera are derived from the same ancestral base about two million years ago. With speciation and evolution, they have acquired many advanced living skills in common, but have also evolved very different living strategies due to different distributions. This thesis is an intensive study of the biology of the mixed-species colonies of these species, the aims of which were to investigate their behavioural relationships and uncover the evolutionary conserved features of their behaviours subsequent to speciation. The results show that the two species can form a stable society to perform normal tasks. First, workers of both species in the mixed-colonies could form the typical retinue behaviour to hetero-species queens, thus indicating that queen pheromones could be spread to and by both species. Secondly, both species did not show significantly different ovarian activation under hetero-species queens, suggesting that the queen pheromones more likely play a role of "honest signal" rather than a "repression" substance in the honeybee colonies. Thirdly, both species could mutually decode each other‘s waggle dances, with unexpectedly low misunderstanding; revealing that the dance language in a dark environment is quite adaptive for cavity-nesting honeybees. Fourthly, workers of both species could cooperate with each other in comb construction, although the combs they built contain many irregular cells. Interestingly, A. cerana workers could be stimulated by A. mellifera workers to perform this task, thus confirming self-organization theory in the colony. Fifthly, A. mellifera workers behaved more "defectively" in thermoregulation, but perhaps because A. cerana workers are more sensitive to changes in hive temperature. Given these differences in strategy, A. mellifera workers‘ performance might in fact reduce conflicts. Lastly, when faced with threats of predatory wasps, both species engaged in aggressive defence. Although they did not learn from each other‘s responses, species-specific strategies were adopted by each of them so that the defence of the mixed-colonies is very effective. I conclude that the two species can adapt to each other‘s efforts and task allocation is reasonably organized allowing mixed-species colonies to reach stability. These results suggest that all of the social behaviours discussed here were highly conserved following speciation. This thesis could provide some clues for the study of honeybee evolution from open-nesting to the transition of cavity-nesting.
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Queens, pseudoqueens and laying workers reproductive competition in the Cape Honeybee (Apis mellifera capensis Eschscholtz)
- Authors: Muerrle, Thomas Martin
- Date: 2008
- Subjects: Honeybee Honeybee -- Reproduction
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5750 , http://hdl.handle.net/10962/d1005437
- Description: In honeybees (Apis mellifera L.) the queen monopolises reproduction. However, especially after queen loss, workers can lay eggs, but are unable to mate. They produce haploid male offspring (drones) from unfertilised eggs via arrhenotokous parthenogenesis. In contrast, workers of the honeybee subspecies Apis mellifera capensis Eschscholtz typically produce diploid female offspring from unfertilised eggs thelytokously. After queen loss and without queen-derived brood A. m. capensis colonies can successfully requeen from worker-derived brood. This, however, is a relatively rare event in wild populations. Moreover, workerderived queens were described to be smaller, more worker-like and reproductively inferior. On the other hand, the fixation of the thelytokous trait relies mainly on sufficient numbers of viable drones produced by worker-derived queens. Small numbers of reproductively inferior worker-derived queens in A. m. capensis populations would be clearly counterintuitive. It is therefore necessary to quantify the significance of worker-dependant queen rearing pathways on the individual (queen) and on population level.Reproductive inferiority of worker-derived queens could not be confirmed on the individual (queen) level when comparing parameters indicating potential reproductive success of queen- and worker-derived queens. Queen- and worker-derived queens clearly showed a congruent range of reproductive performance. In queen rearing preference tests, increased acceptance of worker-derived female larvae was exactly counterbalanced by increased mortality, resulting in an equal number of eclosing virgin queens from an equal number of grafts in both test groups. Larval survival and successful eclosion is a prerequisite for a queen’s reproductive success. I found no difference in eclosion success for queen- and worker-derived virgin queens, indicating a similar potential for reproductive success in both queen types. Assessments of the developmental patterns of colonies headed by both queen and worker-derived queens in long-term experiments revealed no significant differences in reproductive success. Colonies headed by queen-derived queens and colonies headed by worker-derived queens could not be separated when comparing the different developmental pathways observed or from differences in worker-force. Reproductive dominance in A. m. capensis appeared tobe determined by a function of relative compositional and absolute quantitative pheromonal patterns, where individuals, which produce compositionally most queen-like blends in highest quantities, occupy top positions. Queen- and worker-derived virgin queens occupied intermediate positions between pseudoqueens and mated queens. However, no significant differences between the pheromonal status of queen- and worker-derived virgin queens were observed, suggesting a similar range of reproductive dominance for both queen types. In behavioural bioassays queen- and worker-derived virgin queens appeared to be similarly attractive to clustering workers and to drones in a drone congregation area, indicating no differences in potential reproductive success for queens from both origins for those parameters. The significant influence of the queen substance 9-ODA on attractiveness to workers and drones was confirmed. Rare requeening events from worker-derived female brood in queenless A.m. capensis do not satisfactorily explain the fixation of the thelytokous trait at a population level. I observed A. m. capensis worker ovipositing into empty artificial queen cell cups in queen-right colonies. The queen was confined behind a queen excluder grid in a separate compartment of the colony, to imitate reduced pheromonal flow, similar to swarming or superseding colonies. Eggs oviposited by workers in artificial queen cell cups were readily accepted for queen rearing and successful eclosion of viable virgin queens was observed. Consequently I suggested an alternative worker-dependant reproductive pathway in A. m. capensis, which was never described before: In swarming or superseding queenright colonies, laying workers may directly compete with the queen for reproductive success by ovipositing (instead of the queen) into natural queen cell cups. At a population level this reproductive tactic may result in large numbers of worker-derived queens of high reproductive quality in natural populations of A. m. capensis.
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A phenological and bioclimatic analysis of honey yield in South Africa
- Authors: Illgner, Peter Mark
- Date: 2004
- Subjects: Bee culture -- South Africa -- Statistics Honeybee -- South Africa Bee culture -- South Africa -- Management Honey plants -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5813 , http://hdl.handle.net/10962/d1007153
- Description: This study has investigated the interaction between honeybees and their forage plants and the impact of selected climatic variables on honey production in South Africa. Twenty-seven scale-hive records from 25 localities have been used as a measure of colony honey reserves. At least 944 plant species are visited by honeybees in South Africa for their nectar and/or pollen, with more than half providing both rewards. The entire honeybee flora encompasses 532 genera and 137 families. The flowering phenologies of the different reward categories of the indigenous forage plants are all significantly and positively correlated at the 0.05 level. Similarly, species offering both rewards are significantly and positively correlated with the flowering phenology of the null flora. The same results were obtained for correlations between the different reward categories of the exotic forage plants in South Africa. Of the 30 species pairs which fulfilled the criteria for selection, 23 occurred in sympatry, 5 in allopatry and 2 in possible parapatry. There is evidence for both competition and facilitation within different indigenous species pairs. The lack of geographical correlation in the intra-annual variation in honey stores and the near absence of any statistically significant (p < 0.05) honey related intra-annual intracolonial correlations may indicate that the former is more important than the latter for the determination of the level of honey reserves within a colony. Only one statistically significant correlation was found between either scale-hive record from the University of Pretoria Experimental Farm and any of the selected climatic variables. A one month lag period and/or possible seasonal effects were detected for each variable, with the exception of the duration of sunshine, in the autocorrelation analyses. A possible 12 month seasonal period was also identified in the single series fourier analyses for a number of variables. Similarly, 12 months was also the most frequently recurring period in the crossspectral results for the one scale-hive record (H42). Any activities which have an impact on the landscape have the potential to affect honeybees and/or their forage plants. Honeybee crop or plant pollination may also enhance yields for commercial farmers and facilitate rural food security.
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The ecology and control of small hive beetles (Aethina tumida Murray)
- Authors: Ellis, James Douglas
- Date: 2004
- Subjects: Bee culture , Bee culture -- United States , Nitidulidae , Beetles -- Ecology , Beetles -- Control
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5917 , http://hdl.handle.net/10962/d1016253
- Description: The small hive beetle (Aethina tumida Murray) is an endemic scavenger in colonies of honey bee (Apis mellifera L.) subspecies inhabiting sub-Saharan Africa. The beetle only occasionally damages host colonies in its native range and such damage is usually restricted to weakened/diseased colonies or is associated with after absconding events due to behavioral resistance mechanisms of its host. The beetle has recently been introduced into North America and Australia where populations of managed subspecies of European honey bees have proven highly susceptible to beetle depredation. Beetles are able to reproduce in large numbers in European colonies and their larvae weaken colonies by eating honey, pollen, and bee brood. Further, adult and larval defecation is thought to promote the fermentation of honey and large populations of beetles can cause European colonies to abscond, both resulting in additional colony damage. The economic losses attributed to the beetle since its introduction into the United States have been estimated in millions of US dollars. Although beetles feed on foodstuffs found within colonies, experiments in vitro show that they can also complete entire life cycles on fruit. Regardless, they reproduce best on diets of honey, pollen, and bee brood. After feeding, beetle larvae exit the colony and burrow into the ground where they pupate. Neither soil type nor density affects a beetle’s ability to successfully pupate. Instead, successful pupation appears to be closely tied to soil moisture. African subspecies of honey bees employ a complicated scheme of confinement (aggressive behavior toward and guarding of beetles) to limit beetle reproduction in a colony. Despite being confined away from food, adult beetles are able to solicit food and feed from the mouths of their honey bee guards. Remarkably, beetle-naïve European honey bees also confine beetles and this behavior is quantitatively similar to that in African bees. If confinement efforts fail, beetles access the combs where they feed and reproduce. Two modes of beetle oviposition in sealed bee brood have been identified. In the first mode, beetles bite holes in the cappings of cells and oviposit on the pupa contained within. In the second mode, beetles enter empty cells, bite a hole in the wall of the cell, and oviposit on the brood in the adjacent cell. Despite this, African bees detect and remove all of the infected brood (hygienic behavior). Similarly, European bees can detect and remove brood that has been oviposited on by beetles. Enhancing the removal rate of infected brood in European colonies through selective breeding may achieve genetic control of beetles. Additional avenues of control were tested for efficacy against beetles. Reducing colony entrances slowed beetle ingress but the efficacy of this method probably depends on other factors. Further, the mortality of beetle pupae was higher when contacting species of the fungus Aspergillus than when not, making biological control an option. Regardless, no control tested to date proved efficacious at the level needed by beekeepers so an integrated approach to controlling beetles remains preferred. The amalgamation of the data presented in this dissertation contributed to a discussion on the beetle’s ecological niche, ability to impact honey bee colonies in ways never considered, and the ability to predict the beetle’s spread and impact globally.
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Geographical races of the honeybees (Apis Mellifera L.) of the Northern regions of Ethiopia
- Authors: Mohammed, Nuru Adgaba
- Date: 2003
- Subjects: Honeybee -- Ethiopia Honeybee -- Behavior -- Ethiopia Honeybee -- Ethiopia -- Morphology Honeybee -- Ecology -- Ethiopia
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5827 , http://hdl.handle.net/10962/d1007671
- Description: The ideal climatic conditions and diversity of floral resources allow Ethiopia to sustain millions of honeybee colonies. Beekeeping is widely practiced and plays a significant role in the livelihood of the farming community of the country. Despite this, information on the types of geographical races of honeybees, their behavioural characters and the related ecological factors are not established. In this work an attempt was made to characterise the honeybee populations of the northern parts of Ethiopia based on the analyses of morphological, behavioural and ecological characteristics. For morphological analysis, 155 honeybee colony samples from 31 localities were collected. 20 worker honeybees per colony sample totalled 3100 individual worker honeybees and thirteen morphological characters per bee, a total of 40 300 measurements were recorded following Ruttner's (1988) protocols. The behavioural characters such as reproductive swarming, migration, temperament and colony seasonal cycles were assessed based on survey work of a pre-structured questionnaire. Various univariate and multivariate statistical methods were employed to analyse the data. The principal components and step-wise linear discriminant analyses revealed the existence of four discrete morphoclusters or geographical races: A. m. jemenitica from eastern, northwestern and northern arid and semi-arid lowlands, A. m. scutellata from the western humid midlands, A. m. bandasii from sub-moist central highlands and A. m. monticola from the northern mountainous parts of the study areas. These different geographical races were found to be distinctively distributed in the different ecological parts of the study area. High intercolonial and intracolonial variances were observed in all localities, however particularly high values were obtained in areas of transitions between ecological zones indicating a region of gene flow or zones of hybridisation among the statistically defined populations. Such high variances were observed mostly in areas where A. m. jemenitica borders the other geographical races. Moreover. distinct behavioural variations were also noted among these geographical races. Generally. the highland and mountainous honeybees A. m. bandasii and A. m. monticola are relatively larger in body size. have less tendency to swarm. less inclination to migrate and are relatively gentle while the converse traits were noted for the lowlands and midlands honeybees A. m. jemenitica and A. m. scutellata. Both morphological and behavioural variations were highly correlated with environmental factors and the variations seem to be the result of long adaptation to the interrelated ecological factors in their respective areas. Variations in reproductive swarming periods were also noted among these honeybee populations as a result of variations in rainfall pattern. altitude and temperature in their respective ecological areas which are believed to alter the honeybee colonies' seasonal cycles and leads to partial temporal reproductive isolation among these different honeybee populations.
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Multivariate morphometric analysis and behaviour of honeybees (Apis Mellifera L.) in the southern regions of Ethiopia
- Authors: Amssalu, Argaw Bezabeh
- Date: 2003
- Subjects: Honeybee -- Ethiopia -- Behavior Honeybee -- Ethiopia
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5616 , http://hdl.handle.net/10962/d1003130
- Description: Morphometric and behavioural characteristics of honeybees, Apis mellifera were analysed using multivariate and bivariate statistical methods to characterise honeybees of southern Ethiopian region. A total of 33800 morphometric character measurements were taken from 2600 individual worker honeybees of 130 honeybee colonies collected at 26 sampling localities with an average inter-locality distance of 89km to determine the occurrence of morphoclusters. 117 experienced farmer beekeepers and beekeeping experts were interviewed on pre-tested questionnaire to investigate the behavioural characteristics of these honeybees in their respective areas. Morphometric characters associated with pigmentation and body size exhibited a higher discriminant power while forewing venation angles (B4, N23 and 026) lack discriminatory power to segregate honeybees in the southern Ethiopian region. Principal components and discriminant analyses using the most discriminatory morphological characters delineated four statistically distinct morphoclusters in the southern Ethiopian region: the smallest and yellow honeybees, A. m. woyi-gambella which are different from all African honeybees, occur in the western and southern lowlands; the small and yellowiest honeybees, Apis mellifera jemenitica in the eastern escarpment; the largest and darkest honeybees, Apis mellifera bandasii in the central and eastern highlands; and dark honeybees, Apis mellifera scutellata in the wet tropical forests. High intracolonial and intercolonial variances within and between the boundaries of the honeybee groups were detected. The former suggests areas of ecological instability, while the latter areas of transitional or natural hybridisation zones. These areas are characterised by transitional ecological zones having intermediate climate and physiography. Even though honeybees of the southern Ethiopian region are generally dark and small, they show a high tendency to reproductive swarming, migration and aggressiveness, great variation in pigmentation, size and behaviour were observed both within and between the groups. A. m. jemenitica honeybees have a high propensity to migration and less a tendency for reproductive swarming. A. m. bandasii and A. m. scutellata have a high inclination to reproductive swarming and the former has a lesser and the later intermediate propensity to migration. A. m. woyi-gambella honeybees have intermediate swarming and migration tendencies. These results revealed that reproductive swarming and migration are higher in resource-rich and resource-poor areas respectively. Honey plants of the central highlands of Ethiopia are predominantly herbaceous in nature and mainly grow on open and cultivated lands. The bulk of pollen collected came comparatively from few genera. Strong correlation was observed between the intensities of flowering and rainfall. Reproductive swarming and migration occur during high and low intensity of flowering respectively.
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Reproductive conflicts in honeybee colonies
- Authors: Pirk, Christian Walter Werner
- Date: 2002
- Subjects: Honeybee -- Reproduction
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5755 , http://hdl.handle.net/10962/d1005443
- Description: In advanced eusocial hymenopteran societies workers have ovaries and can lay eggs, but are unable to mate. Workers are more related to their own offspring than to every other member of the colony. So worker reproduction contains both worker-worker and worker-queen conflict. The queen- worker conflict is discussed elsewhere, but if the queen mates with more than two males, worker policing should be selected to lower potential conflicts. However in the Cape honeybee it was predicted that worker policing is absent or less expressed than in other honeybee subspecies, because workers produce female offspring thelytokously. So laying workers and their offspring are nearly genetically identical, which results in the fact that other workers are as related to workers derived from eggs laid by the queen as laid by a worker. However, worker reproduction may be costly and therefore worker policing could be an evolutionary adaptation in the Cape honeybee to lower the costs derived from laying worker activity. Indeed, Cape honeybee colonies show efficient egg removal behaviour, suggesting that other factors like colony efficiency could favour egg removal behaviour. Since egg removal behaviour is a colony phenomenon, factors that affect colony performance could also affect egg removal behaviour. Egg removal behaviour was considerably affected by environmental changes, indicating that other tasks have a higher priority than egg removal behaviour. Thousands of queenright colonies of the neighbouring subspecies (A. m. scutellata) were taken over by laying A. m. capensis workers, showing that A. m. capensis workers are facultative social parasites. These observations strongly indicate that laying workers of A. m. capensis are able to evade worker policing and the inhibitory effects of the queen pheromones, but what potential strategies could these laying workers use to increase the survival of their eggs and evade the queen? On the one hand, egg removal behaviour is variable. One behavioural strategy of laying workers to achieve successful reproduction could be that they lay during periods with low egg removal behaviour. On the other hand, the inhibitory effect of the queen’s pheromones diminishes with distance. Maybe the level of egg removal also depends, like the inhibitory effect of the queen pheromones, on the distance from the queen. Indeed, further away from the queen the effect of the queen pheromone and the level of egg removal is reduced, making successful worker reproduction possible. In both subspecies, A. m. capensis and A. m. scutellata, egg removal behaviour is reduced further away from the queen. In the case of A. m. scutellata egg removal is lacking further away from the queen. This explains why colonies of scutellata are so prone to takeovers by laying Cape honeybee workers. One question in the context of parasitic Cape honeybees is how they manage to get into the host colony. One way could be that they get into the colonies during a natural colony merger which is common in African bees. Two unrelated colonies merged and it took them only 24 hours to show effective integration. Because both colonies are unrelated, the potential reproductive conflict among workers should be more strongly expressed than in a normal colony, which is not the result of a merger. Therefore, the effect of nestmate recognition for eggs on the egg removal behaviour was investigated. The results suggest that workers recognise the origin of an egg and that the standard policing experiments overestimate the level of egg removal and only represent relative values. Moreover, the results show that colony specific components on the eggs are more important than a postulated queen egg marking pheromone. Finally, for the first time empirical evidence from a population of the parasitic laying Cape honeybee workers, invading thousands of colonies of A. m. scutellata in northern South Africa, for a short-sighted selection theory is presented.
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The mandibular gland secretions of the Cape honeybee (Apis mellifera capensis ESCH.) : factors affecting the production of the chemical signal and implications for further development of beekeeping in South Africa
- Authors: Jones, Georgina Elizabeth
- Date: 2001
- Subjects: Honeybee Honeybee -- Physiology
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5730 , http://hdl.handle.net/10962/d1005416
- Description: The chemical composition of the mandibular gland extracts of Apis mellifera capensis virgin queens was analysed by gas chromatography-mass spectroscopy. Thirty-seven compounds from various chemical groups including aliphatic and aromatic acids and diacids, phenols, alkanes, amino acids and sugars were identified. Among the identified compounds were the queen mandibular pheromone components 9ODA, 9HDA, HVA and HOB and the other aliphatic acids and phenols considered to be the major components of A.m. capensis mandibular glands. Ontogenetic changes in the concentration of the mandibular gland secretions of virgin queens were largely quantitative in nature with the total volume and that of most of the compounds increasing with queen age. The final level of 9ODA is reached at the premating stage, approximately three days after emergence, when it comprises approximately 87% of the major constituents of the mandibular gland signal. Hostile reactions by workers towards introduced virgin queens can be correlated to the relative proportion of 9ODA present in the mandibular gland secretions. This seems to indicate that it is the complete spectrum of the signal and not individual compounds that determine worker reaction towards introduced queens. Keeping queens singly, with or without workers, in an incubator and in small mating nucleus hives proved to be the most successful methods of queen rearing in respect to survival rate in A.m. capensis. The presence of workers during the ageing of virgin queens was found to significantly affect the chemical composition of the mandibular gland secretions of queens. The reaction of workers towards introduced virgin queens reared under different holding conditions varied, with queens reared with workers eliciting significantly less hostile reactions from workers than those reared without workers. Mated queens from five localities in the Eastern Cape were characterised on the basis of the chemical composition of their mandibular gland secretions and the ratio of 9ODA:10HDA. No significant differences were detected and none of the queens sampled could be considered to be A.m. capensis based on their mandibular gland signal. The findings of this study provide baseline data for the development of a queen-rearing program tailored to the specific requirements of A.m. capensis.
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Wax secretion in the Cape honeybee (Apis mellifera capensis esch.) in relation to juvenile hormone and age polyethism
- Authors: Muller, Nikite Wilhelmine Josephine
- Date: 1993
- Subjects: Honeybee -- Behavior , Honeycombs
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5797 , http://hdl.handle.net/10962/d1005485
- Description: Wax secretion in worker honeybees is significantly related to the age of the worker and, while the secretion pattern remains the same, the absolute amount of wax secreted varies seasonally. Comb building festoons, previously thought to be the site of wax secretion, contain only a fraction of the newly-secreted wax in the nest. Festooning behaviour was also found to be seasonal. The amount of wax secreted by workers was significantly affected by hive. Although age-related changes in behaviour and physiology of worker honeybees appears to be modulated by juvenile hormone (JH), wax secretion is not dependent on JH. Manipulating JH III titres by injecting the hormone and manipulating the only source of the hormone (the corpora allata: CA) did not affect wax secretion. Increasing haemolymph JH titre shortly after ec1osion did not affect the amount of wax produced by workers aged 3 to 21 days, nor could a critical period be found during which elevated hormone titres would affect the rate of wax secretion. Allatectomy of newly eclosed workers did not affect wax production. Removing the putative neural feedback inhibition on the CA did not result in a change in wax production. Implanting CA from older workers into younger workers had no significant effect. Methoprene, a widely-used JH analog, caused reduced wax secretion in workers. It is suggested that methoprene poisons worker honeybees. The results obtained are consistent with an alternative model for wax secretion proposed by Butler (1954). The methodological problems found in this work are present in many other studies. When viewed in this light, the role of JH in polyethism appears dubious and there are alternative models of polyethism that do not have these shortcomings.
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