Giant honeybees (Apis dorsata) mob wasps away from the nest by directed visual patterns

The open nesting behaviour of giant honeybees (Apis dorsata) accounts for the evolution of a series of defence strategies to protect the colonies from predation. In particular, the concerted action of shimmering behaviour is known to effectively confuse and repel predators. In shimmering, bees on the nest surface flip their abdomens in a highly coordinated manner to generate Mexican wave-like patterns. The paper documents a further-going capacity of this kind of collective defence: the visual patterns of shimmering waves align regarding their directional characteristics with the projected flight manoeuvres of the wasps when preying in front of the bees’ nest. The honeybees take here advantage of a threefold asymmetry intrinsic to the prey–predator interaction: (a) the visual patterns of shimmering turn faster than the wasps on their flight path, (b) they “follow” the wasps more persistently (up to 100 ms) than the wasps “follow” the shimmering patterns (up to 40 ms) and (c) the shimmering patterns align with the wasps’ flight in all directions at the same strength, whereas the wasps have some preference for horizontal correspondence. The findings give evidence that shimmering honeybees utilize directional alignment to enforce their repelling power against preying wasps. This phenomenon can be identified as predator driving which is generally associated with mobbing behaviour (particularly known in selfish herds of vertebrate species), which is, until now, not reported in insects.     

Out with the garbage: the parasitic strategy of the mantisfly Plega hagenella mass-infesting colonies of the eusocial bee Melipona subnitida in northeastern Brazil

Between April and June of 2012 mantisflies (Plega hagenella) were found to be extensively parasitizing the nests of two groups of managed colonzies of eusocial stingless bees (Melipona subnitida) in the semi-arid region of northeastern Brazil. The mantisfly larvae developed inside closed brood cells of the bee comb, where each mantispid larva fed on the bee larva or pupa present in a single brood cell. Mature mantispid larvae pupated inside silken cocoons spun in place within their hosts' brood cells then emerged as pharate adults inside the bee colony. Pharate adults were never attacked and killed by host colony workers. Instead, colony workers picked up the pharates and removed them from the nest unharmed, treating them similar to the way that the general refuse is removed from the nest. Adult mantispids subsequently eclosed from their pupal exuviae outside the nest. Manipulative experiments showed that post-eclosion adult mantispids placed back within active bee colonies were quickly attacked and killed. These observations demonstrate that pharate and post-eclosion adults of P. hagenella are perceived differently by colony workers and that delayed adult eclosion is an important functional element in the parasitic life strategy of P. hagenella, allowing adults to escape without injury from the bee colonies they parasitize.     

‘Special agents’ trigger social waves in giant honeybees (Apis dorsata)

Giant honeybees (Apis dorsata) nest in the open and have therefore evolved a variety of defence strategies. Against predatory wasps, they produce highly coordinated Mexican wavelike cascades termed ‘shimmering’, whereby hundreds of bees flip their abdomens upwards. Although it is well known that shimmering commences at distinct spots on the nest surface, it is still unclear how shimmering is generated. In this study, colonies were exposed to living tethered wasps that were moved in front of the experimental nest. Temporal and spatial patterns of shimmering were investigated in and after the presence of the wasp. The numbers and locations of bees that participated in the shimmering were assessed, and those bees that triggered the waves were identified. The findings reveal that the position of identified trigger cohorts did not reflect the experimental path of the tethered wasp. Instead, the trigger centres were primarily arranged in the close periphery of the mouth zone of the nest, around those parts where the main locomotory activity occurs. This favours the ‘special-agents’ hypothesis that suggest that groups of specialized bees initiate the shimmering.     

The ‘I see you’ prey–predator signal of Apis cerana is innate

An ‘I see you’ (ISY) prey–predator signal can co-evolve when such a signal benefits both prey and predator. The prey benefits if, by producing the signal, the predator is likely to break off an attack. The predator benefits if it is informed by the signal that the prey is aware of its presence and can break off what is likely to be an unsuccessful and potentially costly hunt. Because the signal and response co-evolve in two species, the behaviour underlying an ISY signal is expected to have a strong genetic component and cannot be entirely learned. An example of an ISY signal is the ‘shimmering’ behaviour performed by Asian hive bee workers in the presence of their predator Vespa velutina. To test the prediction that bee–hornet signalling is heritable, we let honey bee workers of two species emerge in an incubator so that they had never been exposed to V. velutina. In Apis cerana, the shimmering response developed 48 h post-emergence, was strong after 72 h and increased further over 2 weeks. In contrast, A. mellifera, which has evolved in the absence of Asian hornets, did not produce the shimmering signal. In control tests, A. cerana workers exposed to a non-threatening butterfly did not respond with the shimmering signal.     

A critical number of workers in a honeybee colony triggers investment in reproduction

Social insect colonies, like individual organisms, must decide as they develop how to allocate optimally their resources among survival, growth, and reproduction. Only when colonies reach a certain state do they switch from investing purely in survival and growth to investing also in reproduction. But how do worker bees within a colony detect that their colony has reached the state where it is adaptive to begin investing in reproduction? Previous work has shown that larger honeybee colonies invest more in reproduction (i.e., the production of drones and queens), however, the term ‘larger’ encompasses multiple colony parameters including number of adult workers, size of the nest, amount of brood, and size of the honey stores. These colony parameters were independently increased in this study to test which one(s) would increase a colony’s investment in reproduction via males. This was assayed by measuring the construction of drone comb, the special type of comb in which drones are reared. Only an increase in the number of workers stimulated construction of drone comb. Colonies with over 4,000 workers began building drone comb, independent of the other colony parameters. These results show that attaining a critical number of workers is the key parameter for honeybee colonies to start to shift resources towards reproduction. These findings are relevant to other social systems in which a group’s members must adjust their behavior as a function of the group’s size.     

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     

Scouts behave as streakers in honeybee swarms

Harmonic radar tracking was used to record the flights of scout bees during takeoff and initial flight path of two honeybee swarms. One swarm remained intact and performed a full flight to a destination beyond the range of the harmonic radar, while a second swarm disintegrated within the range of the radar and most of the bees returned to the queen. The initial stretch of the full flight is characterized by accelerating speed, whereas the disintegrating swarm flew steadily at low speed. The two scouts in the swarm displaying full flight performed characteristic flight maneuvers. They flew at high speed when traveling in the direction of their destination and slowed down or returned over short stretches at low speed. Scouts in the disintegrating swarm did not exhibit the same kind of characteristic flight performance. Our data support the streaker bee hypothesis proposing that scout bees guide the swarm by traveling at high speed in the direction of the new nest site for short stretches of flight and slowing down when reversing flight direction.     

Nest odor dynamics in the social wasp Vespula vulgaris

We investigated nest odor dynamics in the common yellow jacket, Vespula vulgaris. In six isolated colonies, we tested the aggression rates toward dead nestmates that had been stored for 10 min, 10 and 19 days outside their colonies at –76 °C. The aggression rate increased from about 12% toward recently killed nestmates up to 30% toward nestmates killed 19 days before the experiment. Obviously, the conserved nest odor profile of the nestmates frozen for several days did not match with that of their colony anymore. This indicates a change of the nest odor within the colony. In a second experiment, we kept two colonies each in one nest box with a complete separation of both neighbor nests by a solid wall inside the box for 28 days. In confrontation experiments, the colony members treated dead foragers from the neighbor nest as aggressively as dead foreign, non-neighbor workers (about 39% each) whereas only about 14% reacted aggressively toward dead nestmates. Seventeen days after the replacement of the solid wall by a metallic grid, which allowed no physical contact but air exchange between the two neighbor colonies, the aggression rates toward foreign workers and nestmates remained relatively unaffected whereas it decreased significantly toward dead neighbors to about 11%. These results suggest a nest odor dynamic caused by volatiles transferred between two adjacent colonies, resulting in an equalization of the former colony specific nest odors. A change of nest odor dynamics influenced by volatiles was so far described only for one ant species at all.     

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.     

The role of moisture in the nest thermoregulation of social wasps

Paper nests of social wasps are intriguing constructions for both, biologists and engineers. We demonstrate that moisture and latent heat significantly influence the thermal performance of the nest construction. Two colonies of the hornet Vespa crabro were investigated in order to clarify the relation of the temperature and the moisture regime inside the nest. Next to fairly stable nest temperatures the hornets maintain a high relative humidity inside the nest. We found that in consequence a partial vapor-pressure gradient between nest and ambient drives a constant vapor flux through the envelope. The vapor flux is limited by the diffusion resistance of the envelope. The driving force of vapor flux is heat, which is consumed through evaporation inside the nest. The colony has to compensate this loss with metabolic heat production in order to maintain a stable nest temperature. However, humidity fluctuations inside the nest induce circadian adsorption and desorption cycles, which stabilize the nest temperature and thus contribute significantly to temperature homeostasis. Our study demonstrates that both mechanisms influence nest thermoregulation and need to be considered to understand the thermodynamic behavior of nests of wasps and social insects in general.     

Bee-hawking by the wasp, <Emphasis Type="BoldItalic">Vespa velutina</Emphasis>, on the honeybees <Emphasis Type="BoldItalic">Apis cerana</Emphasis> and <Emphasis Type="BoldItalic">A</Emphasis>. <Emphasis Type="BoldItalic">mellifera</Emphasis>

The vespine wasps, Vespa velutina, specialise in hawking honeybee foragers returning to their nests. We studied their behaviour in China using native Apis cerana and introduced A. mellifera colonies. When the wasps are hawking, A. cerana recruits threefold more guard bees to stave off predation than A. mellifera. The former also utilises wing shimmering as a visual pattern disruption mechanism, which is not shown by A. mellifera. A. cerana foragers halve the time of normal flight needed to dart into the nest entrance, while A. mellifera actually slows down in sashaying flight manoeuvres. V. velutina preferentially hawks A. mellifera foragers when both A. mellifera and A. cerana occur in the same apiary. The pace of wasp-hawking was highest in mid-summer but the frequency of hawking wasps was three times higher at A. mellifera colonies than at the A. cerana colonies. The wasps were taking A. mellifera foragers at a frequency eightfold greater than A. cerana foragers. The final hawking success rates of the wasps were about three times higher for A. mellifera foragers than for A. cerana. The relative success of native A. cerana over European A. mellifera in thwarting predation by the wasp V. velutina is interpreted as the result of co-evolution between the Asian wasp and honeybee, respectively.     

Chemical compounds of the foraging recruitment pheromone in bumblebees

When the frenzied and irregular food-recruitment dances of bumblebees were first discovered, it was thought that they might represent an evolutionary prototype to the honeybee waggle dance. It later emerged that the primary function of the bumblebee dance was the distribution of an alerting pheromone. Here, we identify the chemical compounds of the bumblebee recruitment pheromone and their behaviour effects. The presence of two monoterpenes and one sesquiterpene (eucalyptol, ocimene and farnesol) in the nest airspace and in the tergal glands increases strongly during foraging. Of these, eucalyptol has the strongest recruitment effect when a bee nest is experimentally exposed to it. Since honeybees use terpenes for marking food sources rather than recruiting foragers inside the nest, this suggests independent evolutionary roots of food recruitment in these two groups of bees.     

Debris removal by head-pushing in A. florea Fabr. honeybees

 The nest of the dwarf honeybee A. florea Fabr. consists of a single comb attached to a tree branch. Recruitment dances take place on the upper surface of the comb that must therefore be kept clear of debris. We report here, for the first time, a behaviour that serves for removing leaves and other foreign objects from the surface of the comb. Individual workers crawl under the object and lift it with their heads, pushing it towards the rim where it eventually slides off the comb. Objects that are heavier or fixed at one end such as leaves are nevertheless lifted and kept away from the surface for up to several minutes. This "head-pushing" is frequently performed without the aid of mandibles, and individuals performing it maintain a distinctive posture, holding the forelegs at an angle without touching the object. Repeated involvement of particular individuals indicate that head-pushers might form a distinct task group. Received: 24 November 1999 / Accepted in revised form: 10 March 2000     

Clonal structure affects the assembling behavior in the Japanese queenless ant <Emphasis Type="Italic">Pristomyrmex punctatus</Emphasis>

The queenless ant Pristomyrmex punctatus (Hymenoptera: Myrmicinae) has a unique society that differs from those of other typical ants. This species does not have a queen, and the workers lay eggs and produce their clones parthenogenetically. However, a colony of these ants does not always comprise members derived from a single clonal line. In this study, we examined whether P. punctatus changes its “assembling behavior” based on colony genetic structure. We prepared two subcolonies—a larger one comprising 200 individuals and a smaller one comprising 100 individuals; these subcolonies were established from a single stock colony. We investigated whether these subcolonies assemble into a single nest. The genetically monomorphic subcolonies (single clonal line) always fused into a single nest; however, the genetically polymorphic subcolonies (multiple clonal lines) did not tend to form a single colony. The present study is the first to demonstrate that the colony genetic structure significantly affects social viscosity in social insects.     

Two experimental tests of the relationship between group stability and aggressive conflict in Polistes wasps

Two field experiments were used to examine how the relative benefits of cooperation influence within-group conflict in foundress associations of the paper wasp Polistes dominulus. P. dominulus foundresses can either nest alone or cooperate with other foundresses. We experimentally manipulated the relative benefits of co-foundress associations vs independent reproduction and tested the effect on aggressive within-group conflict. First, we examined aggression between alpha and beta co-foundresses before and after lower-ranking foundresses were removed. Removal of subordinates increases the relative contributions of the remaining subordinates to group reproductive output as there are fewer adults to care for the brood. Transactional models predict that group conflict over reproductive shares will increase as the relative benefits of grouping increase. As predicted, aggression between the co-foundresses significantly increased following subordinate removal. Second, we experimentally reduced ecological constraints on independent nesting by placing a previously orphaned, adoptable nest comb near the occupied nests. Providing an independent breeding opportunity is predicted to increase the benefits of independent reproduction relative to those of cooperating, thereby reducing group stability and aggression. As predicted, aggression between dominant and subordinate foundresses significantly decreased after the orphaned comb was presented. Therefore, group members sense variation in ecological constraints and relative productivity contributions and quickly modulate their behavior in response. Overall, these two experiments suggest that paper wasps behave as if within-group competition is limited by the threat of group dissolution such that stable groups where cooperation is strongly favored can withstand higher levels of conflict than unstable groups.     

Thorough warm-up before take-off in honey bee swarms

In a bivouacked swarm of honey bees, most individuals are quiescent while a small minority (the scouts) are active in choosing the swarm's future nest site. This study explores the way in which the members of a swarm warm their flight muscles for take-off when the swarm eventually decamps. An infrared camera was used to measure the thoracic (flight muscle) temperatures of individual bees on the surface of a swarm cluster. These are generally the coolest bees in a swarm. The warming of the surface-layer bees occurred mainly in the last 10 min before take-off. By the time a take-off began, 100% of the bees had their flight muscles heated to at least 35°C, which is sufficient to support rapid flight. Take-offs began only a few seconds after all the surface-layer bees had their flight muscles warmed to at least 35°C, but exactly how take-offs are triggered remains a mystery.     

Emissions Scenarios Database and Review of Scenarios

This paper reviews and analyzes more than 400 scenarios of global and regional greenhouse gas emissions and their main driving forces - population, economy, energy intensity, and carbon intensity - drawn from an extensive literature survey and summarized in a database. This new and growing database is available online, which makes summary statistics on these scenarios widely available. The scenarios in the database were collected from almost 200 different literature sources and other scenario evaluation activities. The ultimate objective of the database is to include all relevant global and regional emissions scenarios. This paper shows how the database can be utilized for the analysis of greenhouse gas emissions ranges across the scenarios in the literature and for the analysis of their main driving forces. The scenarios in the database display a large range of future greenhouse gas emissions. Part of the range can be attributed to the different methods and models used to formulate the scenarios, which include simple spreadsheet models, macroeconomic models and systems-engineering models. However, most of the range is due to differences in the input assumptions for the scenarios, in particular of the main scenario driving forces. Special emphasis is given to an analysis of medians and ranges of scenario distributions and the distributions of the main scenario driving forces in the database. The analysis shows that the range for projected population increase in the world, across the scenarios in the database, is the smallest of all main driving forces (about a factor of 3 in 2100). The range of economic growth, measured by the gross world product, and the range of primary energy consumption vary by a factor of 10 in 2100. Carbon intensity of energy, an indicator of the degree of technological change, varies by nearly two orders of magnitude in the year 2100. In addition, this paper presents the first attempt to analyze the relationships among the main scenario driving forces. Subsequent papers in this special issue give further analyses of the relationships among the main scenario driving forces and their other relevant characteristics.     

Do Scaphoideus titanus (Hemiptera: Cicadellidae) nymphs use vibrational communication?

Small Auchenorrhyncha use substrate-borne vibrations to communicate. Although this behaviour is well known in adult leafhoppers, so far no studies have been published on nymphs. Here we checked the occurrence of vibrational communication in Scaphoideus titanus (Hemiptera: Cicadellidae) nymphs as a possible explanation of their aggregative distributions on host plants. We studied possible vibratory emissions of isolated and grouped nymphs, as well as their behavioural responses to vibration stimuli that simulated presence of conspecifics, to disturbance noise, white noise and predator spiders. None of our synthetic stimuli or pre-recorded substrate vibrations from nymphs elicited specific vibration responses and only those due to grooming or mechanical contacts of the insect with the leaf were recorded. Thus, S. titanus nymphs showed to not use species-specific vibrations neither for intra- nor interspecific communication and also did not produce alarm vibrations when facing potential predators. We conclude that their aggregative behaviour is independent from a vibrational communication.     

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.     

Convergent evolution: floral guides, stingless bee nest entrances, and insectivorous pitchers

Several recent hypotheses, including sensory drive and sensory exploitation, suggest that receiver biases may drive selection of biological signals in the context of sexual selection. Here we suggest that a similar mechanism may have led to convergence of patterns in flowers, stingless bee nest entrances, and pitchers of insectivorous plants. A survey of these non-related visual stimuli shows that they share features such as stripes, dark centre, and peripheral dots. Next, we experimentally show that in stingless bees the close-up approach to a flower is guided by dark centre preference. Moreover, in the approach towards their nest entrance, they have a spontaneous preference for entrance patterns containing a dark centre and disrupted ornamentation. Together with existing empirical evidence on the honeybee's and other insects’ orientation to flowers, this suggests that the signal receivers of the natural patterns we examined, mainly Hymenoptera, have spontaneous preferences for radiating stripes, dark centres, and peripheral dots. These receiver biases may have evolved in other behavioural contexts in the ancestors of Hymenoptera, but our findings suggest that they have triggered the convergent evolution of visual stimuli in floral guides, stingless bee nest entrances, and insectivorous pitchers.