Nest Defense Behavior in Colonies from Crosses Between Africanized and European Honey Bees (Apis mellifera L.) (Hymenoptera: Apidae)

Honey bee (Apis mellifera L.) colonies with either European or Africanized queens mated to European or Africanized drones alone or in combination were tested for defensive behavior using a breath test. The most defensive colonies were those with European or Africanized queens mated to Africanized drones. In colonies where both European and Africanized patrilines existed, most of the workers participating in nest defense behavior for the first 30 s after a disturbance were of African patrilines. Nest defense behavior appears to be genetically dominant in honey bees.     

The natural history of nest defence in a stingless bee, Tetragonisca angustula (Latreille) (Hymenoptera: Apidae), with two distinct types of entrance guards

The stingless bee Tetragonsica angustula (Latreille) is the only social bee known that has two different types of nest entrance guards. As in other stingless bees and the honey bee one type stands on, in or near the nest entrance. The second type, so far only known in T. angustula, hovers near the nest entrance. In order to gain further understanding of this unique situation we studied guarding behaviour in both types of guards. Using marked bees, we found that individual worker bees guarded for a long time, up to 20 days, relative to their short, average c. 21 day, lifespan. Relatively few, 33%, individually marked guards were seen performing both types of guarding. The others only acted as standing guards. The bees that did perform both types did so over similar periods of their life. Hovering bouts were 57 min long, interrupted by breaks inside the hive of a few minutes (3.3 ± 1.5 min). Standing bouts were slightly longer (74 min) and also interrupted by short breaks (7.82 ± 6.45 min). Human breath, mimicking a vertebrate intruder, caused the guards to retreat into the nest rather than to attack the intruder. Some colonies protected themselves against intruders by closing the entrance during the night (32% and 56% of colonies during two nights). In summary, our results indicate that nest entrance guarding in T. angustula involves division of labour between the two types, in which most guarding individuals only act as standing guards.     

Major benefits of guarding behavior in subsocial bees: implications for social evolution

Parental care is a behavior that increases the growth and survival of offspring, often at a cost to the parents' own survival and/or future reproduction. In this study, we focused on nest guarding, which is one of the most important types of extended parental care; we studied this behavior in two solitary bee species of the genus Ceratina with social ancestors. We performed the experiment of removing the laying female, who usually guards the nest after completing its provisioning, to test the effects of nest guarding on the offspring survival and nest fate. By dissecting natural nests, we found that Ceratina cucurbitina females always guarded their offspring until the offspring reached adulthood. In addition, the females of this species were able to crawl across the nest partitions and inspect the offspring in the brood cells. In contrast, several Ceratina chalybea females guarded their nests until the offspring reached adulthood, but others closed the nest entrance with a plug and deserted the nest. Nests with a low number of provisioned cells were more likely to be plugged and abandoned than nests with a higher number of cells. The female removal experiment had a significantly negative effect on offspring survival in both species. These nests frequently failed due to the attacks of natural enemies (e.g., ants, chalcidoid wasps, and other competing Ceratina bees). Increased offspring survival is the most important benefit of the guarding strategy. The abandonment of a potentially unsuccessful brood might constitute a benefit of the nest plugging behavior. The facultative nest desertion strategy is a derived behavior in the studied bees and constitutes an example of an evolutionary reduction in the extent of parental care.     

Circadian Activity Rhythm of the Foragers of a Eusocial Bee (Scaptotrigona aff depilis,Hymenoptera, Apidae,Meliponinae) outside the Nest

In all bee colonies of the Meliponinae subfamily, activity inside the nest is temporally organized around the oviposition by the queen, assisted by nurse bees. This class is constituted by young bees that remain inside the nest. In a colony of Scaptotrigona aff depilis, the oviposition cycle occurs in a 3-hour period. The foragers are older bees that collect food for the colony in the field. Other tasks in the nest are performed by workers of ages intermediate between nurses and foragers. With the aim of studying activity rhythms, foragers were kept under constant light, with food constantly available and no flight restriction. The results showed that, although inside the nest the prevailing period is 3 hours, the activity of the foragers is a circadian rhythm, synchronized by the light/dark cycle and probably influenced by other environmental cycles as temperature and the availability of food sources.     

The Fanner Honey Bee: Behavioral Variability and Environmental Cues in Workers Performing a Specialized Task

We tested the hypothesis that Apis mellifera workers exhibit plasticity in moving from fanning to guarding behavior. Bees marked when fanning are more likely to guard than fan on subsequent days, but guard to fanner reversals were common. Our findings suggest that bees can switch between these tasks, but that their bias between the two tasks changes over time, rather than a strict serial progression of worker tasks. The number of fanning workers is positively correlated with ambient temperature and negatively correlated with humidity; this conclusion gives insight into the environmental triggers for worker behavior.     

Age polyethism and defense in a tropical social wasp (Hymenoptera: Vespidae)

Although tapping the nest in simulated vertebrate attack caused all but 1-day-old workers of Polybia occidentalis to rush from inside and cover the envelope, defense itself—attack of a target—was shown to be carried out only by workers older than about 10 days. While foraging was also performed by older workers, the frequency-age distribution of defensive behavior was skewed to a significantly younger age than was that of foraging. The plot of mean individual probability of defending as a function of age was logistic, leveling off at 0.11 by the age of about 13 days. The distribution of the probability of defending among workers 13 days of age was random, indicating that there is no specialized defender class among older workers. Workers that defended were recruited from among both nest workers and foragers. Although foragers were more likely to defend than were nonforagers, nonforagers contributed more individuals to the pool of defenders because they were more numerous.     

Zombie fire ant workers: behavior controlled by decapitating fly parasitoids

Laboratory observations were conducted on four separate red imported fire ant, Solenopsis invicta, colonies that contained workers parasitized by the decapitating fly, Pseudacteon tricuspis. Parasitized S. invicta workers remained inside the nest during parasitoid larval development and left the nest approximately 8 – 10 hours before decapitation by the parasitoid. When parasitized ants left the nest, they were highly mobile, were responsive to tactile stimuli, and showed minimal defensive behavior. Ants ultimately entered into a grass thatch layer, where they were decapitated and the fly maggots pupariated. This study reveals that parasitized ants exhibit behaviors that are consistent with host manipulation to benefit survival of the parasitoid. Received 9 November 2006; revised 26 January 2007; accepted 7 February.     

The role of young guards inXylocopa pubescens

Summary Xylocopa pubescens is a facultatively social species in which two types of guards can be found: 1) old, formerly reproductive guards and 2) young, pre-reproductive guards that usually guard the nest in which they emerged. In this species it is always the dominant female that forages and lays the eggs.This paper focuses on the young females' reasons for guarding. Young guards are characteristically the first females of an emerging brood. They start guarding at an age of 6 days, and continue to do so for on average 10 days, when they start a dominance contest. In comparison to other emerging bees, guards did not receive any more of the incoming food. The presence of young females in the nest was sufficient to deter pollen robbers; the protection was not ameliorated by guarding. Guards did not protect the nest from usurpation by intruding females. The presence of a young guard positively influenced both the number and the duration of foraging flights.All guards attempted to take over dominance inside the nest by fighting with the dominant female. Guards had a probability of 50% of winning this contest, which was distinctly higher than their probability of finding a nest elsewhere. The dominant female was not in all cases the mother of the guard. Therefore, the average increase of indirect fitness by guarding was lower than the expected direct fitness returns from leaving the nest earlier. We therefore conclude that guarding females are environmentally disabled, hopeful reproductives. They may be guarding to determine the right time to risk a fight about dominance and to increase their direct fitness if their attempt to supersede is successful.     

Honeybee (Apis cerana) guards do not discriminate between robbers and reproductive parasites

A hopelessly queenless honeybee colony has only one reproductive option: some workers must produce sons before the colony dies. This requires the workers to curtail egg policing (removal of worker-produced eggs), rendering the colony vulnerable to non-natal reproductive parasitism. In the Western honeybee, Apis mellifera, guarding (prevention of foreign workers from entering a colony) increases in queenless colonies, providing a defence against non-natal parasitism. However, in the closely related Eastern honeybee A. cerana, queenless colonies appear to be more tolerant of bees from other colonies. We presented guards of four A. cerana colonies with three types of workers: nestmate returning foragers, non-nestmate returning foragers and non-nestmates from a laying-worker colony. The latter are likely to have active ovaries, allowing us to test whether guard bees can detect which potential invaders are more likely to be reproductive parasites. After assessing guards’ reactions, we recaptured test bees and dissected them to determine levels of ovary activation. We found that nestmates were accepted significantly more frequently than the other two types of workers. However, there was no difference in the overall acceptance rates of non-nestmate returning foragers and bees from within laying-worker colonies. In addition, ovary-activated workers were no less likely to be accepted than those with inactive ovaries. Interestingly, colonies were more accepting of all three types of test bee after being made queenless. We conclude that, as has been previously suggested, guarding has no specific role in the prevention of non-natal parasitism in A. cerana.     

Observation of group recruitment to prey in a primitive ponerine ant,Amblyopone sp. (reclinata group) (Hymenoptera: Formicidae)

Summary Group recruitment during foraging was observed in a primitive ponerine ant,Amblyopone sp. (reclinata group) under laboratory condition. Workers searched for prey singly; however, if a item of prey was stung by a worker, other ants joined the attack. After the prey became immobile, one of the workers laid a trail directly toward the nest. This scout worker recruited additional workers (between 3 and 33). They formed a single file procession to the point of prey capture, and cooperatively transported the prey. A scout worker could stimulate nest workers to leave the nest without direct contact, and the recruited workers could trace the trail without guidance by the scout worker. This is the first report of recruitment behavior during foraging in the primitive antAmblyopone.     

Ontogeny of the Defensive Stinging Behavior of the Fire Ant, Solenopsis invicta

The fire ant, Solenopsis invicta, appears to deviate from the usual age-related pattern of defensive behavior seen in other social insects; instead of older workers being more defensive than younger ones, they are less so. Here I test this pattern by quantifying changes in the defensive stinging behavior of S. invicta workers of known age. I found defensiveness, measured as both the number of stings delivered and the amount of venom delivered per sting (venom dose), to increase with age initially but then decline after a mid-age peak. This hump-shaped ontogeny may be the result of S. invicta's foraging strategy, which effectively shifts the responsibility of nest defense to workers younger than foraging age. It is S. invicta's mid-aged workers that are the most defensive, probably because they are both physiologically and spatially the most suitable nest defenders.     

German Yellowjacket (Vespula germanica) Foragers Use Odors Inside the Nest to Find Carbohydrate Food Sources

Prior work has shown that yellowjacket waSPS remember food odors and use them as cues when foraging. There is also evidence they have mechanisms to recruit nest mates to highly rewarding food sources, as naïve individuals are more likely to go to food sources with scents similar to those visited by nest mates. We asked whether recruitment requires behavioral stimulation by returning foragers, as in honey bees, or if sampling the food source inside the nest is sufficient. We tested this by eliminating the behavior of returning foragers by inserting a scented sugar solution directly into a Vespula germanica nest. Exiting foragers were given a choice of the test scent and a control scent. WaSPS were more likely to choose the test scent. We conclude that behavioral interactions with returning foragers are not necessary to stimulate nest mates to associate an odor with a food source and search for a resource bearing that odor, and that experience with the scented reward inside the nest is sufficient to achieve this result.     

Construction of Nest Defensive Structure According to Offspring Value and Its Effect on Predator's Attack Decision in Paper Wasps

Various organisms are known to build nests with defensive structures to protect their offspring from predation, but our understanding of plasticity in the nest structure remains poor. In this study, we investigated whether a paper wasp, Polistes chinensis antennalis, adjusted the construction of nest defensive structure according to the value of their offspring, and we also analysed the effect of adjusting the construction of the structure on predator's decision to attack. P. chinensis antennalis foundresses start a colony and maintain her nest alone until the emergence of workers. During this stage, foundresses often construct a defensive structure on cocoon caps of pupae using nest materials (pulp), which prevents predation of pupae by conspecifics from other nests. The value of pupae to the foundress varies among those in a nest, where the value is higher in pupae that spun the cocoon (and initiated pupation) earlier than other pupae in the nest. From field observations, we found that foundresses constructed a larger pulp structure on the cocoons of pupae that cocooned earlier in the order of cocoon spinning, even after considering confounding factors. We also found that the probability of a pupa being attacked by conspecific intruders decreased with the size of pulp structure on the cocoon. This indicates that intruders avoid attacking cocoons with larger pulp structures. Our study indicates that foundresses adjust the construction of nest defensive structures according to their offspring value, and this allows them to protect the high‐value offspring efficiently and effectively.     

Floating and fidelity in nest visitation byCrawfordapis luctuosa (Hymenoptera: Colletidae)

We studied nest visitation behavior of the ground-nesting beeCrawfordapis luctuosa (Colletidae) at the Monteverde Cloud Forest Preserve, Puntarenas Province, Costa Rica. Some females entered many nests, while others consistently entered only one nest. Multiple nest entry was exhibited by bees we called floaters, and those bees that consistently entered only one nest were provisioning bees. Provisioning bees abandoned their old nests after provisioning periods and became nonprovisioning bees that floated over the nesting area. Floating behavior was a nest-searching behavior typified by oscillating flights just above the ground (<15 cm). Floaters investigated available nests and were not observed to interact aggressively with nest owners. Upon establishing a new nest, floaters became provisioning bees.     

An oxytocin/vasopressin-related neuropeptide modulates social foraging behavior in the clonal raider ant

Oxytocin/vasopressin-related neuropeptides are highly conserved and play major roles in regulating social behavior across vertebrates. However, whether their insect orthologue, inotocin, regulates the behavior of social groups remains unknown. Here, we show that in the clonal raider ant Ooceraea biroi, individuals that perform tasks outside the nest have higher levels of inotocin in their brains than individuals of the same age that remain inside the nest. We also show that older ants, which spend more time outside the nest, have higher inotocin levels than younger ants. Inotocin thus correlates with the propensity to perform tasks outside the nest. Additionally, increasing inotocin pharmacologically increases the tendency of ants to leave the nest. However, this effect is contingent on age and social context. Pharmacologically treated older ants have a higher propensity to leave the nest only in the presence of larvae, whereas younger ants seem to do so only in the presence of pupae. Our results suggest that inotocin signaling plays an important role in modulating behaviors that correlate with age, such as social foraging, possibly by modulating behavioral response thresholds to specific social cues. Inotocin signaling thereby likely contributes to behavioral individuality and division of labor in ant societies.

The neuropeptides oxytocin and vasopressin modulate social behavior in vertebrates, but their function in invertebrates is not well understood. Using brain staining and pharmacological manipulations, this study shows that a related neuropeptide, inotocin, affects how ants respond to larvae.     

The influence of the vibration signal on worker interactions with the nest and nest mates in established and newly founded colonies of the honey bee, Apis mellifera

Honey bees adjust cooperative activities to colony needs, based in part on information acquired through interactions with the nest and nest mates. We examined the role of the vibration signal in these interactions by investigating the influence of the signal on the movement rates, cell inspection activity, and trophallaxis behavior of workers in established and newly founded colonies of the honey bee, Apis mellifera. Compared to non-vibrated control bees, vibrated recipients in both colony types exhibited increased movement through the nest and greater cell inspection activity, which potentially increased contact with stimuli that enhanced task performance. Also, compared to controls, recipients in both colony types showed increased rates of trophallactic interactions and spent more time engaged in trophallaxis, which potentially further increased the acquisition of information about colony needs. The vibration signal may therefore help to organize labor in honey bees in part by increasing the rate at which workers obtain information about their colony. Vibrated recipients in the established and newly founded colonies did not differ in any aspect of behavior examined, suggesting that colony developmental state did not influence the degree to which individual workers responded to the signal. However, previous work has demonstrated that newly founded colonies have increased levels of vibration signal behavior. Thus, the vibration signal may help to adjust worker activity to colony conditions partly by stimulating greater numbers of bees to acquire information about colony needs, rather than by altering the level at which individual recipients react to the signal. Received 23 October 2006; revised 15 January 2007; accepted 7 February 2007.     

Respiratory rhythms in stingless bee workers: circadian and ultradian components throughout adult development

The workers of the stingless bee, Melipona quadrifasciata, assume different tasks during their adult life. Newly emerged individuals remain inside the nest, without contact with the external environment. Maturing workers go to more peripheral regions and only the oldest, the foragers, leave the nest. As this diversity of activities implies different metabolic patterns, oxygen consumption has been measured in workers of three different ages: 24–48 h (nurses), 10–15 days (builders), and older than 25 days (foragers). Oxygen consumption of individually isolated workers was determined by intermittent respirometry, under constant darkness and temperature of 25 ± 1°C. Sets of 24-h measurements were obtained from individuals belonging to each of the three worker groups. Rhythmicity has been assessed in the daily (24 h) and ultradian (5–14 h) domains. This experimental design allowed detection of endogenous rhythms without the influence of the social group and without inflicting stress on the individuals, as would be caused by their longer isolation from the colony. Significant 24-h rhythms in oxygen consumption were present in nurses, builders and foragers; therefore, workers are rhythmic from the age of 24–48 h. However, the amplitude of the circadian rhythm changed according to age: nurses showed the lowest values, while foragers consistently presented the largest ones, about ten times larger than the amplitude of nurses’ respiratory rhythm. Ultradian frequencies were detected for all worker groups, the power and frequencies of which varied little with age. This means that the ultradian strength was relatively larger in nurses and apparently maintains some relationship with the queen’s oviposition episodes.     

红火蚁工蚁受侵扰的行为反应强度与蚁巢大小和侵扰强度之间的关系研究

红火蚁Solenopsis invicta Buren的防御行为是其被认定为社会性昆虫的关键特征之一。红火蚁蚁巢受到侵扰时,工蚁会快速涌出搜寻侵扰物并展开攻击。然而,蚁巢受侵扰后工蚁的行为反应与蚁巢大小和侵扰强度之间的关系有待进一步明确。为此,本研究通过野外视频记录并结合室内统计分析的方法来明确不同大小的红火蚁蚁巢受侵扰后工蚁反侵扰行为的动态过程。结果表明,蚁巢大小和受侵扰强度对蚁巢受侵扰后工蚁的反应速度具有显著影响。同等侵扰强度下,与中、大型蚁巢相比,小型蚁巢的工蚁往往表现出更慢的行为反应。尤其在低侵扰强度时,小蚁巢工蚁的反应时间平均0.8 s,而中蚁巢和大蚁巢均仅为0.34 s。当侵扰强度相同时,蚁巢越大,受侵扰后召集工蚁的数量越多;当蚁巢大小相同时,侵扰强度越高,召集工蚁的数量越多。但无论是蚁巢大小还是侵扰强度的不同,红火蚁蚁巢受侵扰后,召集工蚁的数量到达峰值的时间均为30 s左右。研究结果为进一步了解红火蚁的防御行为,科学预防红火蚁的攻击,以及有效开展红火蚁的防治提供了重要的实验依据。     

Bumble bee olfactory information flow and contact-based foraging activation

Nestmate foraging activation and interspecific variation in foraging activation is poorly understood in bumble bees, as compared to honey bees and stingless bees. We therefore investigated olfactory information flow and foraging activation in the New World bumble bee species, Bombus impatiens. We (1) tested the ability of foragers to associate forager-deposited odor marks with rewarding food, (2) determined whether potential foragers will seek out the food odor brought back by a successful forager, and (3) examined the role of intranidal tactile contacts in foraging activation. Bees learned to associate forager-deposited odor marks with rewarding food. They were significantly more attracted to an empty previously rewarding feeder presented at a random position within an array of eight previously non-rewarding feeders. However, foragers did not exhibit overall odor specificity for short-term, daily floral shifts. For two out of three tested scents, activated foragers did not significantly prefer the feeder providing the same scent as that brought back by a successful forager. Finally, bees contacted by the successful forager inside the nest were significantly more likely to leave the nest to forage (38.6% increase in attempts to feed from empty feeders) than were non-contacted bees. This is the first demonstration that tactile contact, a hypothesized evolutionary basal communication mechanism in the social corbiculate bees, is involved in bumble bee foraging activation. Received 4 September 2007; revised 30 May 2008; accepted 15 July 2008.     

Aggressive and Docile Colony Defence Patterns in Apis mellifera. A Retreater–Releaser Concept

Colony defence in Apis mellifera involves a variety of traits ranging from ‘aggressive’ (e.g. entrance guarding, recruitment of flying guards) to ‘docile’ (e.g. retreating into the nest) expression. We tested 11 colonies of three subspecies (capensis, scutellata, carnica) regarding their defensiveness. Each colony was selected as reportedly ‘aggressive’, ‘intermediate’ or ‘docile’ and consisted of about 10,000 bees. We applied three stimulation regimes (mechanical disturbance, exposure to alarm pheromones, and the combination of both) and measured their behaviours by tracking the rates of outflying bees at the entrance sites of the test hives. We provided evidence that for mechanical disturbances the test colonies resolved into two response types, if the ‘immediate’ defence response, assessed in the first minute of stimulation, was taken as a function of foraging: ‘releaser’ colonies allocated flying guards, ‘retreater’ colonies reduced the outside-hive activities. This division was observed irrespective of the subspecies membership and maintained in even roughly changing environmental conditions. However, if pheromone and mechanical stimulation were combined, the variety of colony defensiveness restricted to two further types irrespective of the subspecies membership: six of nine colonies degraded their rate of flying defenders with increasing foraging level, three of the colonies extended their ‘aggressiveness’ by increasing the defender rate with the foraging level. Such ‘super-aggressive’ colonies obviously are able to allocate two separate recruitment pools for foragers and flying defenders.