Vincent Dietemann

Cuticular hydrocarbons mediate discrimination of reproductives and nonreproductives in the ant Myrmecia gulosa

In many species of social insects, the cuticular hydrocarbons of adults vary with both colony identity and individual physiology (oogenesis). Such variations have been shown in some ants and social wasps to function in nestmate recognition, but as yet there is no demonstration of their use by workers to recognize egg layers. We report that in the ant Myrmecia gulosa, workers can discriminate queens and fertile workers from infertile individuals based on distinctive blends of long-chained hydrocarbons present both on the cuticle and in the postpharyngeal gland. The purified hydrocarbon fraction of cuticular extracts from queens elicited high interest in workers, unlike the nonhydrocarbon fraction. However, both fractions were necessary to trigger a response of maximal intensity. In contrast, extracts of mandibular and Dufour glands from queens or infertile workers were not treated differentially by workers. We suggest that cuticular hydrocarbons function as pheromones allowing for recognition of the queen as well as egg-laying workers.

Standard methods for varroa research

Summary Very rapidly after Varroa destructor invaded apiaries of Apis mellifera, the devastating effect of this mite prompted an active research effort to understand and control this parasite. Over a few decades, varroa has spread to most countries exploiting A. mellifera. As a consequence, a large number of teams have worked with this organism, developing a diversity of research methods. Often different approaches have been followed to achieve the same goal. The diversity of methods made the results difficult to compare, thus hindering our understanding of this parasite. In this paper, we provide easy to use protocols for the collection, identification, diagnosis, rearing, breeding, marking and measurement of infestation rates and fertility of V. destructor. We also describe experimental protocols to study orientation and feeding of the mite, to infest colonies or cells and measure the mites susceptibility to acaricides. Where relevant, we describe which mite should be used for bioassays since their behaviour is influenced by their physiological state. We also give a method to determine the damage threshold above which varroa damages colonies. This tool is fundamental to be able to implement integrated control concepts. We have described pros and cons for all methods for the user to know which method to use under which circumstances. These methods could be embraced as standards by the community when designing and performing research on V. destructor.

Miscellaneous standard methods for Apis mellifera research

Summary A variety of methods are used in honey bee research and differ depending on the level at which the research is conducted. On an individual level, the handling of individual honey bees, including the queen, larvae and pupae are required. There are different methods for the immobilising, killing and storing as well as determining individual weight of bees. The precise timing of developmental stages is also an important aspect of sampling individuals for experiments. In order to investigate and manipulate functional processes in honey bees, e.g. memory formation and retrieval and gene expression, microinjection is often used. A method that is used by both researchers and beekeepers is the marking of queens that serves not only to help to locate her during her life, but also enables the dating of queens. Creating multiple queen colonies allows the beekeeper to maintain spare queens, increase brood production or ask questions related to reproduction. On colony level, very useful techniques are the measurement of intra hive mortality using dead bee traps, weighing of full hives, collecting pollen and nectar, and digital monitoring of brood development via location recognition. At the population level, estimation of population density is essential to evaluate the health status and using beelines help to locate wild colonies. These methods, described in this paper, are especially valuable when investigating the effects of pesticide applications, environmental pollution and diseases on colony survival.

Estimating the Density of Honeybee Colonies across Their Natural Range to Fill the Gap in Pollinator Decline Censuses

Although pollinator declines are a global biodiversity threat, the demography of the western honeybee (Apis mellifera) has not been considered by conservationists because it is biased by the activity of beekeepers. To fill this gap in pollinator decline censuses and to provide a broad picture of the current status of honeybees across their natural range, we used microsatellite genetic markers to estimate colony densities and genetic diversity at different locations in Europe, Africa, and central Asia that had different patterns of land use. Genetic diversity and colony densities were highest in South Africa and lowest in Northern Europe and were correlated with mean annual temperature. Confounding factors not related to climate, however, are also likely to influence genetic diversity and colony densities in honeybee populations. Land use showed a significantly negative influence over genetic diversity and the density of honeybee colonies over all sampling locations. In Europe honeybees sampled in nature reserves had genetic diversity and colony densities similar to those sampled in agricultural landscapes, which suggests that the former are not wild but may have come from managed hives. Other results also support this idea: putative wild bees were rare in our European samples, and the mean estimated density of honeybee colonies on the continent closely resembled the reported mean number of managed hives. Current densities of European honeybee populations are in the same range as those found in the adverse climatic conditions of the Kalahari and Saharan deserts, which suggests that beekeeping activities do not compensate for the loss of wild colonies. Our findings highlight the importance of reconsidering the conservation status of honeybees in Europe and of regarding beekeeping not only as a profitable business for producing honey, but also as an essential component of biodiversity conservation.

Varroa destructor : research avenues towards sustainable control

Summary Pollination by honey bees plays a key role in the functioning of ecosystems and optimisation of agricultural yields. Severe honey bee colony losses worldwide have raised concerns about the sustainability of these pollination services. In many cases, bee mortality appears to be the product of many interacting factors, but there is a growing consensus that the ectoparasitic mite Varroa destructor plays the role of the major predisposing liability. We argue that the fight against this mite should be a priority for future honey bee health research. We highlight the lack of efficient control methods currently available against the parasite and discuss the need for new approaches. Gaps in our knowledge of the biology and epidemiology of the mite are identified and a research road map towards sustainable control is drawn. Innovative and challenging approaches are suggested in order to stimulate research efforts and ensure that honey bees will be able to sustainably fulfil their role in the ecosystem.

Do honeybees, Apis mellifera scutellata, regulate humidity in their nest?

Honeybees are highly efficient at regulating the biophysical parameters of their hive according to colony needs. Thermoregulation has been the most extensively studied aspect of nest homeostasis. In contrast, little is known about how humidity is regulated in beehives, if at all. Although high humidity is necessary for brood development, regulation of this parameter by honeybee workers has not yet been demonstrated. In the past, humidity was measured too crudely for a regulation mechanism to be identified. We reassess this issue, using miniaturised data loggers that allow humidity measurements in natural situations and at several places in the nest. We present evidence that workers influence humidity in the hive. However, there are constraints on potential regulation mechanisms because humidity optima may vary in different locations of the nest. Humidity could also depend on variable external factors, such as water availability, which further impair the regulation. Moreover, there are trade-offs with the regulation of temperature and respiratory gas exchanges that can disrupt the establishment of optimal humidity levels. As a result, we argue that workers can only adjust humidity within sub-optimal limits.

Is there a need for conservation of honeybees in Africa

Honeybees are native to Africa and Europe but have been spread worldwide as the basis for an apicultural industry. To date, large and diverse wild populations only remain in Africa. On this continent the beekeeping industry is relatively undeveloped and relies on trapping swarms from wild populations to constitute the managed stocks. Bee breeding is seldom practiced. The situation is therefore different from that of Europe or North America where wild or feral honeybees have almost disappeared and this distinction is important when assessing the conservation status of African honeybees. While African honeybees appear to be more resistant to major diseases, the history of honeybee populations worldwide suggests that their conservation is a necessity. After analyzing the threats to which honeybees are exposed in Africa, we argue that preventive conservation measures are required to maintain the present favorable situation and avoid the declines in populations experienced elsewhere.ZusammenfassungDie Beziehung zwischen Menschen und Honigbienen hat in Afrika verschiedene Formen: In vielen Ländern wird Honigjagd praktiziert; In einigen Ländern ist die traditionelle Bienenhaltung in Körben oder Klotzbeuten weitverbreitet während andere moderne Bienenhaltungstechnologie einsetzen. Der hauptsächliche kontextuelle Unterschied zu anderen Weltregionen, in denen die Honigbienen entweder heimisch sind oder eingeführt wurden, ist die bedeutende Population von freilebenden Bienen, auf denen Honigjagd und Ausbeutung basieren. Im Gegensatz zu Europa machen bewirtschaftete Völker nur einen kleinen Teil der Honigbienenpopulation aus (14–18 aus 310 Mio). Weiterhin basiert die Bienenhaltung in Afrika weitgehend auf dem Fang von wilden Schwärmen und es wird keine oder nur wenig Bienenzucht betrieben. Diese Unterschiede sind wichtig, wenn die Honigbienenpopulationen von den in anderen Teilen der Welt zu beobachtenden Abnahmen geschützt werden sollen. Für Afrika sollten daher besondere politische Maßnahmen entwickelt werden. Auf diesem Kontinent sind die Netzwerke, auf die sich Bienenhalter und die Bienenindustrie stützen können nur sehr schwach entwickelt, und es wurden nur wenige Untersuchungen über die Honigbienen durchgeführt. Es ist daher sehr schwer, genaue Informationen und Zahlen über den Gesundheitszustand der Honigbienenpopulationen zu erhalten und daraus abzuleiten, ob Schutzmaßnahmen notwendig sind. Angesichts der Größe der Wildpopulation und dem Fehlen von Berichten über Epidemien könnte man schließen, dass die Wildpopulation in Afrika nicht gefährdet ist und daher von den Imkern ausgebeutet werden kann. Obwohl afrikanische Honigbienen den meisten Krankheiten ausgesetzt sind, die weltweit Honigbienen schädigen und sie diesen widerstehen können, sind der Verlust von Habitaten und Bejagung zwei Faktoren, denen europäische Bienen in geringerem Ausmaß ausgesetzt sind (da freilebende Honigbienen praktisch nicht mehr vorkommen und die bewirtschafteten Völker in vom Menschen hergestellten Beuten leben). Diese und andere noch nicht identifizierte Faktoren (z. B. neue Krankheiten) können Honigbienen negativ beeinflussen und ihre Wirkung könnte unter Umständen zu Verlusten in ähnlichem Ausmaß führen, wie sie andernorts beobachtet werden. Um zu verhindern, dass etwas Ähnliches in Afrika passieren kann, schlagen wir die Schaffung und Verbesserung sowie eine effektive Inkraftsetzung von Regulationen vor, die auf eine Erhaltung der afrikanischen Honigbienen abzielen. Da Afrika das Ursprungsland von Apis mellifera darstellt und das einzige Weltgebiet, in dem eine große Wildpopulation erhalten ist, sind Anstrengungen zur Erhaltung nicht nur zum Erhalt von einzigartigen Populationen vonnöten, sondern auch, um das Funktionieren des Ökosystems und der landwirtschaftlichen Produktion sicherzustellen, die beide von den Bestäubungsleistungen der Honigbienen abhängig sind.

Seasonal prevalence of pathogens and parasites in the savannah honeybee (Apis mellifera scutellata)

The loss of Apis mellifera L. colonies in recent years has, in many regions of the world, been alarmingly high. No single cause has been identified for these losses, but the interactions between several factors (mostly pathogens and parasites) have been held responsible. Work in the Americas on honeybees originating mainly from South Africa indicates that Africanised honeybees are less affected by the interplay of pathogens and parasites. However, little is known about the health status of South African honeybees (A. m. scutellata and A. m. capensis) in relation to pathogens and parasites. We therefore compared the seasonal prevalence of honeybee pathogens (viruses, bacteria, fungi) and parasites (mites, bee lice, wax moth, small hive beetles, A. m. capensis social parasites) between sedentary and migratory A. m. scutellata apiaries situated in the Gauteng region of South Africa. No significant differences were found in the prevalence of pathogens and parasites between sedentary and migratory apiaries. Three (Black queen cell virus, Varroa destructor virus 1 and Israeli acute paralysis virus) of the eight viruses screened were detected, a remarkable difference compared to European honeybees. Even though no bacterial pathogens were detected, Nosema apis and Chalkbrood were confirmed. All of the honeybee parasites were found in the majority of the apiaries with the most common parasite being the Varroa mite. In spite of hosting few pathogens, yet most parasites, A. m. scutellata colonies appeared to be healthy.

Hormonal correlates of reproductive status in the queenless ponerine ant, Streblognathus peetersi

In colonies of the queenless ant Streblognathus peetersi, dominance interactions produce a reproductive hierarchy in which one individual, the alpha, is capable of producing offspring while her subordinates remain infertile. Based on differences between behaviour and cuticular hydrocarbon profiles, the subordinates can be further divided into high and low ranking workers. Although it had been shown previously that alphas treated with a juvenile hormone analog lose their reproductive status, little was known of the endocrinological basis of dominance in this species. To elucidate the underlying endocrinology of these three ranks, we measured the individual in vitro rate of juvenile hormone (JH) production of excised corpora allata, and the ecdysteroid titer of pooled hemolymph samples. Production of JH was highest in low-ranking workers, intermediate in high rankers, and almost undetectable in alphas. Ecdysteroid titers were low for low rankers, but were more than twice as high for both high rankers and alphas. The results support the hypothesis that JH suppresses ovarian function in these queenless ants, and suggest that ecdysteroids may be responsible for stimulating vitellogenin production. The possible role of these hormones as behavioural modulators is also discussed.

Individual versus social pathway to honeybee worker reproduction (Apis mellifera): pollen or jelly as protein source for oogenesis?

Honeybee workers, Apis mellifera, can reproduce in queenless colonies. The production of queen-like pheromones may be associated with their reproductive activity and induce nestmates to respond by feeding them. Such frequent trophallaxis could supply their protein needs for oogenesis, constituting a social pathway to worker reproduction. However, some individuals can develop ovaries without producing queen pheromones. The consumption of protein-rich pollen could be an alternative solitary pathway for them to satisfy this dietary requirement. In order to investigate the way in which workers obtain proteins for oogenesis, we created orphaned worker groups and determined ovarian and pheromonal development in relation to pollen consumption of selected workers. Individuals that did not consume pollen had significantly more developed ovaries and produced significantly more queen mandibular pheromone than workers that fed directly on pollen. Our results suggest that workers producing queen-like secretions are fed trophallactically. However, reproductive workers that lacked queen pheromones had consumed little or no pollen, suggesting that they also obtained trophallaxis. Although pollen consumption might contribute to sustaining oogenesis, it does not appear to be sufficient. Trophallaxis as a means of obtaining proteins seems to be necessary to attain reproductive status in queenless honeybee colonies.