Vibration Signal Behavior of Waggle-dancers in Swarms of the Honey Bee,Apis mellifera (Hymenoptera: Apidae)

We hypothesize two functions of the vibration signal (dorsal ventral abdominal vibration = DVAV) during swarming in honey bees: 1. it enhances recruitment to the specific sites advertised by the waggle dancers which also perform the vibration signal; and 2. it acts as a nonspecific modulatory signal to stimulate activity in other bees. The stimulation of activity invoked by the second hypothesis might include increasing nest-site scouting and dance following early in the house-hunting process or rousing quiescent bees to prepare them for lift-off late in the process, or both. In studies of neotropical African bee swarms in Costa Rica and European bees in California we tested these hypotheses by looking for associations between production of vibration signals by nest-site recruiters and site attractiveness (indicated by which site was ultimately chosen and by distance from the swarm since swarms may have a distance preference). Overall, bees dancing for the chosen sites performed vibration signals to the same extent as those dancing for the other sites. There were no distance differences between sites whose scouts did and did not vibrate other bees. These results are inconsistent with the hypothesis that the vibration signal enhances recruitment to especially high quality sites and they support the hypothesis that it plays a general excitatory role in the context of house hunting by swarming bees.     

Age and Behavior of Honey Bee Workers,Apis mellifera,that Interact with Drones

Colony reproduction in honey bees involves complex interactions between sterile workers and reproductive castes. Although worker–queen interactions have been studied in detail, worker–drone interactions are less well understood. We investigated caste interactions in honey bees by determining the age and behavior of workers that perform vibration signals, trophallaxis, and grooming with drones. Workers of all ages could engage in the different interactions monitored, although workers that performed vibration signals on drones were significantly older than those engaging in trophallaxis and grooming. Only 3–8% of workers engaged in the different behaviors were monitored. Compared with workers that performed vibration signals only on workers (‘worker vibrators’), those that performed signals on both workers and drones (‘drone vibrators’) had greater movement rates inside the nest, higher vibration signaling rates, and were more likely to have an immediate association with foraging. Both worker vibrators and drone vibrators contacted drones of all ages as they moved through the nest. However, drone vibrators contacted drones at higher rates, contacted slightly, but significantly younger drones, and were more likely to engage in trophallaxis and grooming with drones, in addition to vibrating them. Taken together, our results suggest that tiny proportions of workers belonging to separate, but overlapping age groups provide most of the care received by adult drones, and that drone vibrators comprise a subset of signalers within a colony that have an increased tendency to contact and interact with drones. Vibratory, tactile signals are involved in colony reproductive and movement decisions in a number of species of bees, wasps and ants, and may provide valuable tools for investigating caste interactions in many insect societies.     

Distinguishing signals and cues: bumblebees use general footprints to generate adaptive behaviour at flowers and nest

Chemicals used in communication are divided into signals and cues. Signals are moulded by natural selection to carry specific meanings in specific contexts. Cues, on the other hand, have not been moulded by natural selection to carry specific information for intended receivers. Distinguishing between these two modes of information transfer is difficult when animals do not perform obvious secretion behaviours. Although a number of insects have been suspected of leaving cues at food sites and nest entrances, studies have not attempted to experimentally distinguish between cues and signals. Here, we examine the chemical composition of the scent marks left by the bumblebee Bombus terrestris at food sites and compare it to those found at a neutral location. If bees are depositing a cue, we expect the same chemicals to be found at both sites, but if they deposit a signal we only expect to find the scent marks at the food site. We were also interested in identifying the chemicals left at the nest entrance to determine if they differed from those used to mark food sites. We find that bees deposit the same chemicals at food, nest and neutral sites. Therefore, bumblebees leave behind general chemical footprints everywhere they walk and we propose that they learn to use these footprints in a manner that ultimately enhances their fitness, for example, to improve their foraging efficiency and locate their nest. Experimentally, distinguishing these two modes of information transfer is crucial for understanding how they interact to shape animal behaviour and what chemical bouquets are under natural selection. Handling Editor: Heikki Hokkanen     

Age and Behavior of Honey Bees,Apis mellifera (Hymenoptera: Apidae), that Perform Vibration Signals on Workers

The vibration signal is one of the most commonly occurring communication displays in honey bee (Apis mellifera) colonies. It may function in a ‘modulatory’ manner, because it causes a nonspecific increase in activity that enhances a variety of behaviors depending upon the age and caste of the recipient. We examined honey bee workers that performed vibration signals on other workers in three observation hives, each containing a population of marked bees of known age. In all three colonies, the mean age of the first performance of the vibration signal was significantly different from the mean age at which workers first performed waggle dances, carried pollen loads, or attended the queen. However, workers of all ages, except those less than 3 d old, could perform vibration signals. In older workers of foraging age, signal performance was most closely associated with recent foraging success. Younger workers that vibrated did not appear to be early-maturing foragers and thus their signals were probably not influenced by food collection. Rather, for these preforaging-age workers, signal performance was associated more with periods of orientation flight, during which younger bees learn the location of the nest and surrounding landmarks. Thus, the vibration signal may be triggered by different stimuli in different worker age classes. Because it elicits a general increase in activity in all recipients, the signal may help adjust many different colony behaviors simultancously to changes in foraging success and colony development.     

Contextual modulation of visuomotor associations in bumble-bees (Bombus terrestris)

We performed the following experiment to investigate whether contextual cues can prevent interference during the acquisition of potentially competing visuomotor associations. In the bumble-bee (Bombus terrestris). Bees discriminated between horizontally and vertically orientated gratings of black and white stripes in order to reach a feeder and between different diagonally orientated gratings to gain access to their nest. Once bees were well trained on these two tasks, the discrimination task at the nest was changed so that bees had to distinguish between horizontal and vertical gratings at both sites. Whereas they still approached the horizontal grating to reach food, they now had to approach a vertical grating in order to return to their nest. The new task was learnt rapidly. Errors at the feeder did not increase during or after the acquisition of this potentially competing behaviour at the nest entrance. For a brief period during acquisition, bees showed some hesitation at the feeder and hovered for slightly longer before choosing between horizontals and verticals. After acquisition was complete, bees showed a slight increase in their preference for verticals over a more ambiguous stimulus of an array of dots. These findings are consistent with the hypothesis that different contextual signals are associated with approaching the nest or approaching the feeder, and that these contextual signals facilitate learnt associations between orientation detectors and motor commands.     

The Role of the Vibration Signal during Nest‐Site Selection by Honey Bee Swarms

The vibration signal may influence nest‐site selection by honey bee swarms by enhancing scouting and recruitment. We investigated this hypothesis by comparing (1) the number of nest sites and the distances communicated by nest‐site dancers on swarms from which vibrators were and were not removed and (2) the behavior of scouts visiting higher‐quality (HQ) and lower‐quality (LQ) nest boxes. The removal of vibrators from swarms did not alter the number of nest sites investigated, the distances traveled to nest sites, or the time required to select a new nest cavity. In contrast, vibrator removal tripled the time required for swarms to achieve liftoff after a cavity had been selected, although all swarm eventually became airborne and moved to a new site. About 14% of the scouts that visited the HQ and LQ nest boxes performed vibration signals; however, nest‐box quality did not influence the tendency to produce the signal or intermix vibration signals and recruitment dances. However, we did find a significant, positive correlation between overall levels of vibration signal activity and nest‐site recruitment during the house‐hunting process. When viewed in concert, our results suggest that the vibration signal contributes to the house‐hunting process by operating in a non‐specific manner that may enhance scouting and recruitment in general during nest‐site selection and facilitate rapid swarm liftoff after a new nest site has been chosen. The vibration signal is therefore a component in the cascade of communication signals that orchestrate house‐hunting and colony relocation decisions.     

Seasonal spatial dynamics and causes of nest movement in colonies of the invasive Argentine ant (Linepithema humile)

Abstract.  1. Colony organisation and movement behaviour of the Argentine ant ( Linepithema humile ) was studied over 3 years in field populations in California and in captive colonies in the laboratory. This invasive species is highly polydomous and unicolonial; colonies consist of expansive and fluid networks of nests and trails. The spatial and temporal organisation of colonies may contribute to ecological dominance.
2. Argentine ant nests and inter-nest trails shift in size, abundance, and location, so that colony networks are spatially contracted in the winter and expanded spring to autumn. Colonies occupy permanent sites; ants migrated to and from the same winter nest locations year after year, and occupied 30% of the same nests repeatedly during seasonal migrations.
3. Nests were moved on average 2–3 m. Forty-two per cent were occupied less than 1 month, 4% the entire study, and the other 54% lasted 3.9 ± 2.3 months (mean ± SD).
4. Nests were located within 2–4 m of woody plants, in warm sites in the winter and cool sites in the summer. Both humidity and food availability influenced nest-site choice in laboratory colonies. However, when faced with a trade-off between factors, the ants chose humid nest boxes over nest boxes near food, and ants moved nests only in response to changes in humidity and not distance to food.
5. The results indicate that L. humile colonies are seasonally polydomous, and that nest movements are driven by changes in microclimate. Colony organisation maintains high local density and increases food supply, which may improve the competitive ability of L. humile colonies and reduce opportunities for species coexistence.     

Mating Behavior in Halictine Bees (Hymenoptera: Halictidae)

Studied two types of mate-orientation behavior in ♂♂ of the sweat bee, Lasioglossum rohweri: microterritoriality and patrolling. Observations were made in both the laboratory and the field. These ♂♂ behave uniquely for Hymenoptera in that 2–5 of them establish microterritories immediately around the same nest entrance. Some ♂♂ are exclusively patrollers, flying about flowers and nest sites. Rendezvous places (locations where animals are likely to find mates) in bees are: flowers, nesting sites, vegetative parts of plants, nest entrances, air, and nests. Territoriality in male bees may have arisen independently in 7 of the 9 families of bees.     

Post-fledging behaviour in Golden Eagles Aquila chrysaetos: onset of juvenile dispersal and progressive distancing from the nest

Thirteen juvenile Golden Eagles Aquila chrysaetos were tracked during their first year of life using satellite telemetry. Distances to the nest attained during that period and the age at the onset of juvenile dispersal were explored. The performance of nine different criteria to determine that age was analysed. In general, after a brief period of restricted movements around the nest, the average distance to the nest increased with time. Maximum distances to the nest ranged between 57.7 and 184.3 km, and were considerably greater in females (mean ± sd, 138.5 ± 44.5 km) than in males (70.5 ± 14.0 km). No sex difference was observed in the age at which that distance was attained (males: 329 ± 32 days, females: 312 ± 20 days). The onset of juvenile dispersal took place around the fifth month of life (September in Spain). Eight of the nine criteria provided similar results, suggesting that in Spain dispersal starts when birds are between 140 and 180 days old, and that the post-nestling period lasts between 60 and 120 days. For future studies, to determine the age at which the onset of juvenile dispersal occurs, we recommend the use of either the first day on which individuals were located beyond the mean distance between nests of different pairs (10 km in our study area), or the date of the record midway between the first and the last location recorded during the month in which the maximum variability in the distance to the nest was observed.     

Bumble bees (Bombus terrestris) store both food and information in honeypots

Social insect foragers often transmit information about foodsources to nest mates. In bumble bees (Bombus terrestris), forexample, successful foragers use excited motor displays anda pheromone as communication signals. In addition, bees couldmake use of an indirect pathway of information flow, via thehoney stores. We show here that, indeed, bees in the nest continuouslymonitor honeypots and sample their contents, thus obtaininginformation on supply and demand of nectar. When there is aninflux of nectar into the nest, the colony deploys more workersfor foraging. The number of new foragers depends on sugar concentration.Foragers returning with high-quality sugar solution displaymore "excited runs" on the nest structure. The recruits' response,however, does not depend on modulated behavior by foragers:more workers start to forage with high quality of incoming nectar,even when this nectar is brought by a pipette. Moreover, weshow that the readiness of bees to respond to recruitment signalsor incoming nectar also depends on colony demand. When colonynectar stores are full, the response of bees to equal amountsof nectar influx is smaller than when stores are empty. Whencolony nectar stores are depleted, foragers spend more timerunning excitedly and less time probing pots in the nest andrun with higher average speed, possibly to disperse the alertingpheromone more efficiently. However, more bees respond to nectarinflux to empty stores, whether or not this is accompanied byforager signals. Thus, honeypots serve to store informationas well as food.     

Nesting aggregation as a predictor of brood parasitism in mason bees (Osmia spp.)

1. Parasitism can be an important source of mortality for insect populations; however, we know little about the factors influencing vulnerability of wild bees to parasites. Mason bees (genus Osmia; Hymenoptera: Megachilidae) are important pollinators of crops and wild plants and are vulnerable to attack by brood parasites. High nest densities may increase rates of brood parasitism by attracting disproportionate numbers of parasites. 2. Three years of field observations from multiple sites were analysed to assess whether mason bee brood parasitism increased with host density. Mason bees were allowed to nest in artificial nesting blocks and establish natural variation in nesting density. Nest cells constructed by bees were checked for the presence of parasite eggs. 3. Parasitism of nest cells strongly increased with the number of actively nesting bees at a nesting block. Mason bees showed no preference for nesting in blocks that were occupied or unoccupied by other mason bees. Parasitism also increased with the number of days a nest was provisioned and decreased over the course of the season. Nest cells constructed last in a nest were significantly more parasitised than inner cells, despite being sealed against invasions. 4. These findings show positively density‐dependent parasitism in mason bees. They also suggest that bees terminate parasitised nests, causing parasitised cells to become outermost nest cells – a behaviour that may represent a defence against parasites. Our results have implications for the management of mason bees as agricultural pollinators, as cultivating them at high densities could reduce offspring survival.     

Age polyethism in the leaf-cutting ant Acromyrmex subterraneus brunneus Forel, 1911 (Hym., Formicidae)

Abstract  This study describes and quantifies the behavioural acts of two laboratory colonies of Acromyrmex subterraneus brunneus by investigating worker age polyethism. Twenty-nine behavioural acts were recorded during the 19-week observation period. Young individuals performed tasks inside the nest related to brood care and care for the fungus garden, whereas older individuals performed activities outside the nest such as foraging and activities in the waste chamber. The average longevity (±SD) was 108.21 ± 3.30, 109.15 ± 1.92 and 122.71 ± 1.55 days for large, medium and small workers, respectively. The small-sized workers presented a higher probability of reaching older age than large- and medium-sized workers. This study describes task switching according to age polyethism and the relationship of physical and temporal subcastes.     

The Use of the Vibration Signal and Worker Piping to Influence Queen Behavior during Swarming in Honey Bees, Apis mellifera

We investigated worker regulation of queen activity during reproductive swarming by examining the rates at which workers performed vibration signals and piping on queens during the different stages of the swarming process. Worker–queen interactions were first examined inside observation hives during the 2–3 wk that preceded the issue of the swarm (pre‐swarming period) and then inside the swarm clusters during the period that preceded liftoff and relocation to a new nest site (post‐swarming period). Queen court size did not differ between the pre‐ and post‐swarming periods, but workers fed the queens less inside the swarm clusters. Workers performed vibration signals on the queens at increasing rates throughout the pre‐swarming period inside the natal nest, but rarely or never vibrated the queen inside the swarm. Piping was performed on the queens during both the pre‐ and post‐swarming periods and always reached a peak immediately before queen flight. During the final 2–4 h before swarm liftoff, queens were increasingly contacted by waggle dancers for nest sites, some of which piped the queen. The vibration signal may operate in a modulatory manner to gradually prepare the queen for flight from the natal nest, and the cumulative effects of the signal during the pre‐swarming period may make further vibrations on the queen unnecessary when inside the swarm cluster. In contrast, worker piping may function in a more immediate manner to trigger queen takeoff during both the pre‐ and post‐swarming periods. Workers that vibrate and pipe the queen tend to be older, foraging‐age bees. The regulation of queen activity during colony reproduction may therefore be controlled largely by workers that normally have little contact with queens, but help to formulate colony reproductive and movement decisions.     

Segregation of temporal and spatial distribution between kleptoparasites and parasitoids of the eusocial sweat bee, Lasioglossum malachurum (Hymenoptera: Halictidae, Mutillidae)

Cuckoo bees and velvet ants use different resources of their shared host bees, the former laying eggs on the host pollen stores and the latter on immature stages. We studied the activity patterns of the cuckoo bee Sphecodes monilicornis and the velvet ant Myrmilla capitata at two nesting sites of their host, the social digger bee Lasioglossum malachurum , over a 3 year period. Due to the difference in host exploitation, we expected different temporal patterns of the two natural enemies as well as a positive spatial association with host nest density for both species. At a daily level, S. monilicornis was more abundant between 10.00 and 15.00 h, while M. capitata was most active in the early morning and late afternoon; both species activities were independent from host provisioning activity. The activity of cuckoo bees was in general positively correlated with the density of open host nests (but not with the total number of nests), while that of velvet ants was rarely correlated with this factor. Sphecodes monilicornis was seen both attacking the guard bees and directly entering into the host nests or digging close to nest entrances, while M. capitata only gained access to host nests through digging. We conclude that the temporal and spatial segregation between the two species may be, at least partially, explained both by the different resources exploited and by the different dynamics of host interactions.     

Age and Behavior of Honey Bees,Apis mellifera (Hymenoptera: Apidae), that Perform Vibration Signals on Queens and Queen Cells

The vibration signal of the honey bee (Apis mellifera) may play a central role in the regulation of queen behavior during reproductive swarming and supersedure. We examined honey bee workers that performed vibration signals on queens and developing queen cells in three observation hives, each containing a population of marked bees of known age. In all three colonies, workers of all ages greater than 2 d old could perform vibration signals on queens and queen cells. However, most signals were performed by a small proportion of the bees of greater than 10 d of age. Relatively few workers less than 10 d old vibrated queens and queen cells, even though this age-group is typically associated with queen care. Thus, the regulation of queen behavior by the vibration signal may occur primarily through a relatively small subset of older workers that, under most circumstances, have only limited involvement with queens. It is unclear what triggers the vibrating of queens. Workers producing vibration signals did not differ from same-age non-vibrating controls in rate of locomotion in the hive or in task performance, and they rarely engaged in foraging, even though the majority of observed bees were of foraging age; vibrators also did not spend more time with queens and queen cells compared with controls. Vibration signals performed on queens and queen cells therefore do not appear to be influenced by task performance or increased contact with queens.     

Individual bumblebees vary in response to disturbance: a test of the defensive reserve hypothesis

Bees may leave their nest in the event of an attack, but this is not their only response. Here, we examine the behavior of those individuals that remain inside the nest during a disturbance. Specifically, we test the hypothesis that bee workers usually exhibiting high levels of inactivity (i.e., ‘lazy’ bees) may function as defensive reserves that are more likely to respond when the colony is disturbed. We explore this hypothesis by simulating vertebrate attacks by vibrating or blowing carbon dioxide into two colonies on alternating days and measuring the movements and tasks performed by bees inside the nest. Our results show that regardless of the disturbance type, workers increase guarding behavior after a disturbance stops. Although previously inactive bees increased their movement speed inside the nest when the disturbance was vibration, they were not more likely to leave the nest (presumably to attack the simulated attacker) or switch to guarding behavior for any disturbance type. We therefore reject the hypothesis that inactive Bombus impatiens bumblebees act as defensive reserves, and propose alternative hypotheses regarding why many workers remain inactive inside the nest.     

‘Bee Hotels’ as Tools for Native Pollinator Conservation: A Premature Verdict?

Society is increasingly concerned with declining wild bee populations. Although most bees nest in the ground, considerable effort has centered on installing ‘bee hotels’—also known as nest boxes or trap nests—which artificially aggregate nest sites of above ground nesting bees. Campaigns to ‘save the bees’ often promote these devices despite the absence of data indicating they have a positive effect. From a survey of almost 600 bee hotels set up over a period of three years in Toronto, Canada, introduced bees nested at 32.9% of sites and represented 24.6% of more than 27,000 total bees and wasps recorded (47.1% of all bees recorded). Native bees were parasitized more than introduced bees and females of introduced bee species provisioned nests with significantly more female larva each year. Native wasps were significantly more abundant than both native and introduced bees and occupied almost 3/4 of all bee hotels each year; further, introduced wasps were the only group to significantly increase in relative abundance year over year. More research is needed to elucidate the potential pitfalls and benefits of using bee hotels in the conservation and population dynamics of wild native bees.     

Standard yet unusual mechanisms of long-distance dispersal: seed dispersal of Corymbia torelliana by bees

Mellitochory, seed dispersal by bees, has been implicated in long-distance dispersal of the tropical rain forest tree, Corymbia torelliana (Myrtaceae). We examined natural and introduced populations of C. torelliana for 4 years to determine the species of bees that disperse seeds, and the extent and distance of seed dispersal. The mechanism of seed dispersal by bees was also investigated, including fruit traits that promote dispersal, foraging behaviour of bees at fruits, and the fate of seeds. The fruit structure of C. torelliana , with seed presented in a resin reward, is a unique trait that promotes seed dispersal by bees and often results in long-distance dispersal. We discovered that a guild of four species of stingless bees, Trigona carbonaria, T. clypearis, T. sapiens , and T. hockingsi, dispersed seeds of C. torelliana in its natural range. More than half of the nests found within 250 m of fruiting trees had evidence of seed transport. Seeds were transported minimum distances of 20–220 m by bees. Approximately 88% of seeds were dispersed by gravity but almost all fruits retained one or two seeds embedded in resin for bee dispersal. Bee foraging for resin peaked immediately after fruit opening and corresponded to a peak of seed dispersal at the hive. There were strong correlations between numbers of seeds brought in and taken out of each hive by bees ( r =  0.753–0.992, P  < 0.05), and germination rates were 95 ± 5%. These results showed that bee-transported seeds were effectively dispersed outside of the hive soon after release from fruits. Seed dispersal by bees is a non-standard dispersal mechanism for C. torelliana, as most seeds are dispersed by gravity before bees can enter fruits. However, many C. torelliana seeds are dispersed by bees, since seeds are retained in almost all fruits, and all of these are dispersed by bees.     

Influence of rough handling on Osmia lignaria (Hymenoptera: Megachilidae) nest establishment in commercial orchards

Osmia lignaria Say (Hymenoptera: Megachilidae) can be used to pollinate fruit trees. Populations are sometimes difficult to sustain because some female bees fail to establish at provided nesting sites. We address the hypothesis that rough handling of overwintered O. lignaria results in decreased establishment. We tested this by shaking (200 rpm for 2 min) overwintering bees as a proxy for rough handling. Bees were then released in an orchard, and nest establishment of shaken and unshaken bees was recorded. There was no significant difference in the proportion of shaken and unshaken females that nested, indicating that rough handling of overwintering bees does not discourage nest establishment.     

Applying geographic profiling used in the field of criminology for predicting the nest locations of bumble bees

We tested whether geographic profiling (GP) can predict multiple nest locations of bumble bees. GP was originally developed in the field of criminology for predicting the area where an offender most likely resides on the basis of the actual crime sites and the predefined probability of crime interaction. The predefined probability of crime interaction in the GP model depends on the distance of a site from an offender's residence. We applied GP for predicting nest locations, assuming that foraging and nest sites were the crime sites and the offenders’ residences, respectively. We identified the foraging and nest sites of the invasive species Bombus terrestris in 2004, 2005, and 2006. We fitted GP model coefficients to the field data of the foraging and nest sites, and used GP with the fitting coefficients. GP succeeded in predicting about 10-30% of actual nests. Sensitivity analysis showed that the predictability of the GP model mainly depended on the coefficient value of buffer zone, the distance at the mode of the foraging probability. GP will be able to predict the nest locations of bumble bees in other area by using the fitting coefficient values measured in this study. It will be possible to further improve the predictability of the GP model by considering food site preference and nest density.