Prison construction and guarding behaviour by European honeybees is dependent on inmate small hive beetle density

Increasing small hive beetle (Aethina tumida Murray) density changes prison construction and guarding behaviour in European honeybees (Apis mellifera L.). These changes include more guard bees per imprisoned beetle and the construction of more beetle prisons at the higher beetle density. Despite this, the number of beetles per prison (inmate density) did not change. Beetles solicited food more actively at the higher density and at night. In response, guard bees increased their aggressive behaviour towards beetle prisoners but did not feed beetles more at the higher density. Only 5% of all beetles were found among the combs at the low density but this percentage increased five-fold at the higher one. Successful comb infiltration (and thus reproduction) by beetles is a possible explanation for the significant damage beetles cause to European honeybee colonies in the USA.     

Social encapsulation of beetle parasites by Cape honeybee colonies (Apis mellifera capensis Esch.)

Worker honeybees (Apis mellifera capensis) encapsulate the small hive beetle (Aethina tumida), a nest parasite, in propolis (tree resin collected by the bees). The encapsulation process lasts 1-4 days and the bees have a sophisticated guarding strategy for limiting the escape of beetles during encapsulation. Some encapsulated beetles died (4.9%) and a few escaped (1.6%). Encapsulation has probably evolved because the small hive beetle cannot easily be killed by the bees due to its hard exoskeleton and defensive behaviour.     

Small hive beetles survive in honeybee prisons by behavioural mimicry

We report the results of a simple experiment to determine whether honeybees feed their small hive beetle nest parasites. Honeybees incarcerate the beetles in cells constructed of plant resins and continually guard them. The longevity of incarcerated beetles greatly exceeds their metabolic reserves. We show that survival of small hive beetles derives from behavioural mimicry by which the beetles induce the bees to feed them trophallactically. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at htpp://dx.doi.org/10.1007/s00114-002-0326-y.     

The alternative Pharaoh approach: stingless bees mummify beetle parasites alive

Workers from social insect colonies use different defence strategies to combat invaders. Nevertheless, some parasitic species are able to bypass colony defences. In particular, some beetle nest invaders cannot be killed or removed by workers of social bees, thus creating the need for alternative social defence strategies to ensure colony survival. Here we show, using diagnostic radioentomology, that stingless bee workers (Trigona carbonaria) immediately mummify invading adult small hive beetles (Aethina tumida) alive by coating them with a mixture of resin, wax and mud, thereby preventing severe damage to the colony. In sharp contrast to the responses of honeybee and bumblebee colonies, the rapid live mummification strategy of T. carbonaria effectively prevents beetle advancements and removes their ability to reproduce. The convergent evolution of mummification in stingless bees and encapsulation in honeybees is another striking example of co-evolution between insect societies and their parasites.     

Hot spots in the bee hive

Honeybee colonies (Apis mellifera) maintain temperatures of 35-36°C in their brood nest because the brood needs high and constant temperature conditions for optimal development. We show that incubation of the brood at the level of individual honeybees is done by worker bees performing a particular and not yet specified behaviour: such bees raise the brood temperature by pressing their warm thoraces firmly onto caps under which the pupae develop. The bees stay motionless in a characteristic posture and have significantly higher thoracic temperatures than bees not assuming this posture in the brood area. The surface of the brood caps against which warm bees had pressed their thorax were up to 3.2°C warmer than the surrounding area, confirming that effective thermal transfer had taken place.     

Does the Spatial Distribution of the Parasitic Mite Varroa jacobsoni Oud. (Mesostigmata: Varroidae) in Worker Brood of Honey Bee Apis Mellifera L. (Hymenoptera: Apidae) Rely on an Aggregative Process?

  Varroa jacobsoni is an ectoparasite of honey bees which reproduces in capped brood cells. Multi-infestation is frequently observed in worker brood and can be interpreted as an aggregative phenomenon. The aim of this study was to determine whether the distribution of V. jacobsoni in worker brood cells relies on a random or an aggregative process. We studied the distribution of Varroa females in capped worker brood at similar age by comparing, by a Monte Carlo test, the observed frequency distribution of mites per cell to simulated distributions based on a random process. A complementary approach, using the "nearest neighbor distances" (NND) with Monte Carlo tests, was investigated to study the spatial distribution (a) between mites in different cells and (b) between infested cells in brood. The observed distributions did not differ significantly from that expected by a random process, and we conclude that there is no aggregation during invasion of V. jacobsoni in worker brood. Received: 29 April 1999 / Accepted in revised form: 26 August 1999     

Endemic pollinator response to organic vs. conventional farming and landscape context in the Cape Floristic Region biodiversity hotspot

The effects of organic management and landscape context on two highly endemic and important pollinator taxa (bees and monkey beetles) were studied in the Cape Floristic Region (CFR) biodiversity hotspot, where a quarter of the land is intensively managed mostly for grape production. This functional group was chosen as there is concern worldwide over pollinator decline, particularly in regions of high levels of endemism of both flora and fauna such as the CFR. Species richness and abundance of bees and monkey beetles were sampled and compared between organic and conventional vineyards, with natural vegetation as reference habitat. Species richness of monkey beetles, but not the species richness of bees, benefited from organic compared to conventional vineyard management. Our findings support the hypothesis that the effects of landscape context and management may be taxon dependent. Monkey beetles have been suggested as strong indicators of disturbance. These beetles are among the most important pollinator guilds in the CFR, as they help to sustain the endemic vegetation of this region. Protection of the remaining natural habitat in close proximity to farmland is an imperative conservation strategy in this region.     

Stenusine,an antimicrobial agent in the rove beetle genus <Emphasis Type="Italic">Stenus</Emphasis> (Coleoptera,Staphylinidae)

Stenusine is well known as the alkaloid, discharged by the rove beetle, genus Stenus Latreille (Coleoptera, Staphylinidae). The Stenus beetles employ the alkaloid as an escape mechanism when on water surfaces. In the case of danger, they lower their abdomen and emit stenusine from their pygidial glands. Stenusine shows a low surface tension and therefore a high spreading pressure; these properties propel the beetle quickly over the water. Many Steninae do not live in habitats with open waters, but in detritus, leaf litter, mosses, etc. This raises the possibility that stenusine might also have another function, e.g., as antibiotic or fungicide. Stenus beetles show an intense grooming behaviour. With gas chromatography-mass spectrometry analyses we could prove that they cover themselves with their secretion. To tests its antimicrobial properties we conducted agar diffusion tests with stenusine and norstenusine, another substance that is abundant in most Stenus species. Both compounds have an antimicrobial effect on entomopathogenic bacteria and fungi. Stenusine not only allows for an extraordinary method of locomotion on water surfaces, it also protects the Steninae from being infested with microorganisms.     

Caps and gaps: a computer model for studies on brood incubation strategies in honeybees (<Emphasis Type="Italic">Apis mellifera carnica</Emphasis>)

In addition to heat production on the comb surface, honeybee workers frequently visit open cells (“gaps”) that are scattered throughout the sealed brood area, and enter them to incubate adjacent brood cells. We examined the efficiency of this heating strategy under different environmental conditions and for gap proportions from 0 to 50%. For gap proportions from 4 to 10%, which are common to healthy colonies, we find a significant reduction in the incubation time per brood cell to maintain the correct temperature. The savings make up 18 to 37% of the time, which would be required for this task in completely sealed brood areas without any gaps. For unnatural high proportions of gaps (>20%), which may be the result of inbreeding or indicate a poor condition of the colony, brood nest thermoregulation becomes less efficient, and the incubation time per brood cell has to increase to maintain breeding temperature. Although the presence of gaps is not essential to maintain an optimal brood nest temperature, a small number of gaps make heating more economical by reducing the time and energy that must be spent on this vital task. As the benefit depends on the availability, spatial distribution and usage of gaps by the bees, further studies need to show the extent to which these results apply to real colonies. M. Fehler and M. Kleinhenz contributed equally to this work.     

Ecosystem engineering by leaf-rolling mites enhances arthropod diversity

Bombardier beetles (Coleoptera, Carabidae, Brachininae) possess a remarkable defense mechanism where a hot chemical spray is released from the tip of their abdomen, with an audible explosive sound. To date, the repellent properties of these chemicals have been tested against a limited number of taxa, such as amphibians and insects. To investigate the impact of bombardier beetle defenses on avian predators, feeding trials were conducted using the bombardier beetle (Pheropsophus jessoensis) and the Japanese quail (Coturnix japonica), a sympatric and generalist predator. All naïve, hand-reared quail attacked live beetles, indicating the absence of an innate aversion to them. However, most of the quail rejected consuming the beetles whether or not the beetles sprayed them with chemicals. Naïve quail also rejected dead P. jessoensis individuals. These results support the recent hypothesis that it is not essential for P. jessoensis to spray noxious chemicals to deter predators. We also found that some of the quail exposed to live P. jessoensis remembered to avoid them for up to 5 weeks. Our results provide the first evidence of the repelling effects of bombardier beetle defense mechanisms on avian predators.

     

Fever in honeybee colonies

 Honeybees, Apis spp., maintain elevated temperatures inside their nests to accelerate brood development and to facilitate defense against predators. We present an additional defensive function of elevating nest temperature: honeybees generate a brood-comb fever in response to colonial infection by the heat-sensitive pathogen Ascosphaera apis. This response occurs before larvae are killed, suggesting that either honeybee workers detect the infection before symptoms are visible, or that larvae communicate the ingestion of the pathogen. This response is a striking example of convergent evolution between this "superorganism" and other fever-producing animals. Received: 2 September 1999 / Accepted in revised form: 28 February 2000     

Heat Shielding: A Novel Method of Colonial Thermoregulation in Honey Bees

 Honey bees, Apis mellifera, maintain constant colony temperatures throughout the year. Honey bees fan their wings to cool the colony, and often spread fluid in conjunction with this behavior to induce evaporative cooling. We present an additional, previously undescribed mechanism used by the honey bee to maintain constant colony temperature in response to localized temperature increases. Worker bees shield the comb from external heat sources by positioning themselves on hot interior regions of the hive's walls. Although honey comb and brood comb were both shielded, the temperature-sensitive brood received a greater number of heat shielders and was thus better protected from overheating. Heat shielding appears to be a context-dependent adaptive behavior performed by worker bees who would previously have been considered "unemployed." Received: 16 November 1998 / Accepted in revised form: 31 March 1999     

Firewalls in bee nests—survival value of propolis walls of wild Cape honeybee (<Emphasis Type="Italic">Apis mellifera capensis</Emphasis>)

The Cape bee is endemic to the winter rainfall region of South Africa where fires are an integral part of the ecology of the fynbos (heathland) vegetation. Of the 37 wild nests in pristine Peninsula Sandstone Fynbos in the Cape Point section of Table Mountain National Park that have been analyzed so far, only 22 could be accessed sufficiently to determine the existence of a propolis wall of which 68% had propolis walls which entirely enclosed their openings. The analysis of the 37 wild nests revealed that 78% occurred under boulders or in clefts within rocks, 11% in the ground, 8% in tree cavities, and 3% within shrubs. The analysis of 17 of these nests following a fire within the park revealed that the propolis walls materially protected the nests and retarded the fire with all the colonies surviving. The bees responded to the smoke by imbibing honey and retreating to the furthest recess of their nest cavity. The bees were required to utilize this honey for about 3 weeks after which fire-loving plants appeared and began flowering. Considerable resources were utilized in the construction of the propolis walls, which ranged in thickness from 1.5 to 40 mm (mean 5 mm). Its physical environment determines the nesting behavior of the Cape bee. The prolific use of propolis serves to insulate the nest from extremes of temperature and humidity, restricts entry, camouflages the nest, and acts as an effective fire barrier protecting nests established mostly under rocks in vegetation subjected to periodic fires.     

Individual lifetime pollen and nectar foraging preferences in bumble bees

Foraging specialization plays an important role in the ability of social insects to efficiently allocate labor. However, relatively little is known about the degree to which individual bumble bees specialize on collecting nectar or pollen, when such preferences manifest, and if individuals can alter their foraging preferences in response to changes in the colony workforce. Using Bombus impatiens, we monitored all foraging visits made by every bee in multiple colonies and showed that individual foragers exhibit consistent lifetime foraging preferences. Based upon the distribution of foraging preferences, we defined three forager types (pollen specialists, nectar specialists, and generalists). In unmanipulated colonies, 16–36?% of individuals specialized (≥90?% of visits) on nectar or pollen only. On its first day of foraging, an individual’s foraging choices (nectar only, pollen only, or nectar and pollen) significantly predicted its lifetime foraging preferences. Foragers that only collected pollen on their first day of foraging made 1.61- to 1.67-fold more lifetime pollen foraging visits (as a proportion of total trips) than foragers that only collected nectar on their first foraging day. Foragers were significantly larger than bees that stayed only in the nest. We also determined the effect of removing pollen specialists at early (brood present) or later (brood absent) stages in colony life. These results suggest that generalists can alter their foraging preferences in response to the loss of a small subset of foragers. Thus, bumble bees exhibit individual lifetime foraging preferences that are established early in life, but generalists may be able to adapt to colony needs.     

Pesticide exposure in honey bees results in increased levels of the gut pathogen <Emphasis Type="Italic">Nosema</Emphasis>

Global pollinator declines have been attributed to habitat destruction, pesticide use, and climate change or some combination of these factors, and managed honey bees, Apis mellifera, are part of worldwide pollinator declines. Here we exposed honey bee colonies during three brood generations to sub-lethal doses of a widely used pesticide, imidacloprid, and then subsequently challenged newly emerged bees with the gut parasite, Nosema spp. The pesticide dosages used were below levels demonstrated to cause effects on longevity or foraging in adult honey bees. Nosema infections increased significantly in the bees from pesticide-treated hives when compared to bees from control hives demonstrating an indirect effect of pesticides on pathogen growth in honey bees. We clearly demonstrate an increase in pathogen growth within individual bees reared in colonies exposed to one of the most widely used pesticides worldwide, imidacloprid, at below levels considered harmful to bees. The finding that individual bees with undetectable levels of the target pesticide, after being reared in a sub-lethal pesticide environment within the colony, had higher Nosema is significant. Interactions between pesticides and pathogens could be a major contributor to increased mortality of honey bee colonies, including colony collapse disorder, and other pollinator declines worldwide.     

Experience,but not distance,influences the recruitment precision in the stingless bee <Emphasis Type="Italic">Scaptotrigona mexicana</Emphasis>

Recruitment precision, i.e. the proportion of recruits that reach an advertised food source, is a crucial adaptation of social bees to their environment. Studies with honeybees showed that recruitment precision is not a fixed feature, but it may be enhanced by factors like experience and distance. However, little is known regarding the recruitment precision of stingless bees. Hence, in this study, we examined the effects of experience and spatial distance on the precision of the food communication system of the stingless bee Scaptotrigona mexicana. We conducted the experiments by training bees to a three-dimensional artificial patch at several distances from the colony. We recorded the choices of individual recruited foragers, either being newcomers (foragers without experience with the advertised food source) or experienced (foragers that had previously visited the feeder). We found that the average precision of newcomers (95.6 ± 2.61%) was significantly higher than that of experienced bees (80.2 ± 1.12%). While this might seem counter-intuitive on first sight, this “loss” of precision can be explained by the tendency of experienced recruits to explore nearby areas to find new rewarding food sources after they had initially learned the exact location of the food source. Increasing the distance from the colony had no significant effect on the precision of the foraging bees. Thus, our data show that experience, but not the distance of the food source, affected the patch precision of S. mexicana foragers.     

Flower-visiting behavior of male bees is triggered by nectar-feeding insects

Bees are important pollinators for many flowering plants. Female bees are thought to be more effective pollinators than male bees because they carry much more pollen than males. Males of some solitary bee species are known to patrol near flowers that females visit. Because patrolling males visit flowers to mate or defend their territories, they may function as pollinators. However, the significance of patrolling males to pollination has not been studied. We studied males of a solitary bee, Heriades fulvohispidus (Megachilidae), patrolling near flowers and visiting flowers that attracted nectar-feeding insects, including conspecifics, on the Ogasawara (Bonin) Islands. To test the hypothesis that patrolling male bees may function as pollen vectors, we compared the frequency of visits by H. fulvohispidus to flowers of an endemic plant, Schima mertensiana (Theaceae); comparisons were made among flowers with a dead H. fulvohispidus, a dead beetle, a piece of plastic, and nothing (control flowers). Patrolling H. fulvohispidus more frequently visited flowers with a dead conspecific, a dead beetle, or a piece of plastic than the control flowers. Our experiment demonstrates that nectar-feeding insects (including conspecifics and other insects) enhance the flower-visiting frequency of patrolling H. fulvohispidus males on S. mertensiana flowers. Furthermore, we observed S. mertensiana pollen on patrolling males as well as females, suggesting that male bees may also function as pollen vectors.     

First evidence of regular common cuckoo, <Emphasis Type="Italic">Cuculus canorus</Emphasis>, parasitism on eastern olivaceous warblers, <Emphasis Type="Italic">Hippolais pallida elaeica</Emphasis>

Coevolution is defined as specialized relationships between species that lead to a reciprocal evolutionary change. A particularly suitable model system for studying coevolution is the interactions between obligate avian brood parasites and their hosts. The common cuckoo (Cuculus canorus, hereafter cuckoo) is a well-known brood parasite, which utilizes a range of smaller passerines as hosts. However, warblers of the genus Hippolais have rarely been reported as being victims of cuckoos, and furthermore, few data exist on the occurrence of antiparasite defenses in these hosts. In this study, we examined possible host–parasite coevolution between cuckoos and eastern olivaceous warblers (Hippolais pallida elaeica, hereafter olivaceous warblers) in three closely situated areas in northwestern Bulgaria. The olivaceous warbler has never been reported to be a regular cuckoo host. However, the present study, carried out in 2001–2003 shows that the olivaceous warbler is regularly and heavily parasitized by the cuckoo in this area. Parasitism rate was high (26.6%, 34/128) and consistent among years, with some variation between areas. The cuckoo egg mimicry was moderately good, and olivaceous warbler rejection rate of such eggs was 50%. Cuckoo eggs laid in olivaceous warbler nests had a whitish to whitish-green ground color, and the majority appeared to be distinctly different from cuckoo eggs found in other host species in the area. The olivaceous warbler proved to be a rather good host for cuckoos as 20.6% (7/34) of cuckoo eggs laid produced fledglings, a breeding success comparable to other suitable hosts in Europe. This is the first in-depth study of brood parasitism in a warbler of the genus Hippolais, and cuckoos parasitizing olivaceous warblers probably represent a previously unknown gens.     

Gigantism in honeybees: Apis cerana queens reared in mixed-species colonies

The development of animals depends on both genetic and environmental effects to a varying extent. Their relative influences can be evaluated in the social insects by raising the intracolonial diversity to an extreme in nests consisting of workers from more than one species. In this study, we studied the effects of mixed honeybee colonies of Apis mellifera and Apis cerana on the rearing of grafted queen larvae of A. cerana. A. mellifera sealed worker brood was introduced into A. cerana colonies and on emergence, the adults were accepted. Then, A. cerana larvae were grafted for queen rearing into two of these mixed-species colonies. Similarly, A. cerana larvae and A. mellifera larvae were also grafted conspecifically as controls. The success rate of A. cerana queen rearing in the test colonies was 64.5%, surpassing all previous attempts at interspecific queen rearing. After emergence, all virgin queens obtained from the three groups (N=90) were measured morphometrically. The A. cerana queens from the mixed-species colonies differed significantly in size and pigmentation from the A. cerana control queens and closely approximated the A. mellifera queens. It is inferred that these changes in the A. cerana queens reared in the mixed-species colonies can be attributed to feeding by heterospecific nurse bees and/or chemical differences in royal jelly. Our data show a strong impact of environment on the development of queens. The results further suggest that in honeybees the cues for brood recognition can be learned by heterospecific workers after eclosion, thereby providing a novel analogy to slave making in ants.     

Predatory behavior in a necrophagous bee<Emphasis Type="Italic"> Trigona hypogea</Emphasis> (Hymenoptera; Apidae,Meliponini)

Although most bees feed on nectar and pollen, several exceptions have been reported. The strangest of all is the habit found in some neotropical stingless bees, which have completely replaced pollen-eating by eating animal protein from corpses. For more than 20 years, it was believed that carrion was the only protein source for these bees. We report that these bees feed not only off dead animals, but on the living brood of social wasps and possibly other similar sources. Using well developed prey location and foraging behaviors, necrophagous bees discover recently abandoned wasps nests and, within a few hours, prey upon all immatures found there.