Effect of "spraying" by fighting honey bee queens (Apis mellifera L.) on the temporal structure of fights

To reproduce, social insect colonies rear sexual progeny, and young queens start a new colony either without (independently) or with the help of workers (dependently). When colony reproduction is dependent and young queens are produced in excess, conflicts are predicted to occur. Honey bee colonies reproduce dependently by swarming. The mother queen leaves with a Óprime swarmÓ before daughter queens reach adulthood. More queens are produced than can obtain sufficient worker force, and emerging queens often fight to death. Surviving queen(s) inherit the established nest or a portion of workers which then depart in an ÓafterswarmÓ. Honey bee queens show numerous adaptations for fighting and conflict with other queens, such as early venom production and fast development. During fights one queen often releases rectal fluid. The function of this ÓsprayingÓ behaviour is unclear. Previous studies have reported that it can both attract and repel workers, and observations that it also interrupts fights. Possible functions of spraying are to affect worker intervention in fights, act as a chemical weapon, or lower worker attention towards contaminated queens. We staged fights between 24 queen pairs to investigate the temporal pattern of behaviour in spraying and non-spraying fights. Spraying occurred in 67% of the fights. In the majority of spraying fights (87%) it occurred upon physical contact. Spraying fights were characterized by significantly lower proportion of escalated aggression and a significantly shorter first escalated bout. This provides quantitative evidence that spraying interrupts fights and suggests that its function is to directly provide a temporary respite to queens

Characterization of queen specific components of the fluid released by fighting honey bee queens

Swarming honey bee (Apis mellifera L.) colonies rear supernumerary young queens that compete for the limited resources (workers) necessary for founding a new colony. Young queens often fight to death. Queens show several adaptations to fight and conflict, such as short developmental time and early onset of venom production. During fights, queens often release rectal fluid with a strong smell of wine grapes, after which they temporarily stop fighting. This potentially reduces individual overall risk of deadly injury. The fluid and one of its components, ortho-aminoacetophenone, were previously found to have a pheromonal effect on workers, but the evidence is equivocal. Recently, it has been suggested that the effects of this substance may be context- or concentration-specific. We performed semi-quantitative gas-chromatography mass spectrometry (GC/MS) analysis of the fluid (1) released by queens during their first fight, (2) released during a subsequent fight, and (3) obtained by dissecting the hindgut of queens and (4) of workers. Following preliminary results by Page et al. 1988 (Experientia 44: 270-271), we scored presence / absence of eight substances. Five substances (ortho-aminoacetophenone, decanoic acid, dodecanoic acid, octyl decanoate, and decyl decanoate) were characteristic of queens only. ortho-Aminoacetophenone was detected in all queen and in none of the worker samples, in agreement with previous findings that worker faeces do not have any pheromonal effect. The fluid released by queens on their second fight also contained ortho-aminoacetophenone, but in smaller quantities. These data substantiate previous bioassay results, and provide estimates of ortho-aminoacetophenone concentration as required to design experiments addressing the function and adaptive significance of this behaviour

Working-class royalty: bees beat the caste system

The struggle among social classes or castes is well known in humans. Here, we show that caste inequality similarly affects societies of ants, bees and wasps, where castes are morphologically distinct and workers have greatly reduced reproductive potential compared with queens. In social insects, an individual normally has no control over its own fate, whether queen or worker, as this is socially determined during rearing. Here, for the first time, we quantify a strategy for overcoming social control. In the stingless bee Schwarziana quadripunctata, some individuals reared in worker cells avoid a worker fate by developing into fully functional dwarf queens

Correlated expression of phenotypic and extended phenotypic traits across stingless bee species: worker eye morphology, foraging behaviour, and nest entrance architecture.

Abstract: Stingless bees are the most species-rich group of eusocial bees and show great diversity in behaviour, ecology, nest architecture, colony size, and worker morphology. How this variation relates to varying selection pressures and constraints is not well understood. Variation can be caused by selection acting on behavioural or morphological traits, both alone and in correlation across traits. Here we tested whether behavioural and morphological traits important for foraging and defence are linked to nest-entrance architecture, an extended phenotype relevant to both foraging and nest defence. Using 23 species we investigated whether eye size, nest entrance size, landing behaviour and foraging method show cross-species correlations. A phylogenetically-controlled comparative analysis revealed that species with relatively smaller eyes build relatively larger entrances, which in turn are associated with faster landing approaches and fewer landing errors by foragers, both of which could reduce predation risk. Concerning foraging, mass-recruiting species have c. 10-times larger entrance holes than species with a solitary foraging strategy. Larger entrances could help species with mass recruitment to rapidly increase forager traffic or mount a strong defensive response when under attack. Our results show that studying correlations among different traits helps understand phenotypic diversity in species rich groups

Bourgeois behavior and freeloading in the colonial orb web spider Parawixia bistriata (Araneae, Araneidae).

Spiders of the tropical American colonial orb weaver Parawixia bistriata form a communal bivouac in daytime. At sunset, they leave the bivouac and construct individual, defended webs within a large, communally built scaffolding of permanent, thick silk lines between trees and bushes. Once spiders started building a web, they repelled other spiders walking on nearby scaffolding with a bounce behavior. In nearly all cases (93%), this resulted in the intruder leaving without a fight, akin to the bourgeois strategy, in which residents win and intruders retreat without escalated contests. However, a few spiders (6.5%) did not build a web due to lack of available space.Webless spiders were less likely to leave when bounced (only 42% left) and instead attempted to freeload, awaiting the capture of prey items in nearby webs. Our simple model shows that webless spiders should change their strategy from bourgeois to freeloading satellite as potential web sites become increasingly occupied

Effects of group composition and level of selection in the evolution of cooperation in artificial ants

Since ants and other social insects have long generation time, it is very difficult for biologists to study the origin of complex social organization by guided evolution (a process where the evolution of a trait can be followed during experimental evolution). Here we use colonies of artificial ants implemented as small mobile robots with simple vision and communication abilities to explore these issues. In this paper, we present results concerning the role of relatedness (genetic similarity) and levels of selection (individual and colony-level selection) on the evolution of cooperation and division of labor in simulated ant colonies. In order to ensure thorough statistical analysis, the evolutionary experiments, herein reported, have been carried out using "minimalist" simulations of the collective robotics evolutionary setup. The results show that altruistic behaviors have low probability of emerging in heterogeneous colonies evolving under individual-level selection and that colonies with high genetic relatedness display better performance

Anarchy in the UK: Detailed genetic analysis of worker reproduction in a naturally occurring British anarchistic honeybee, Apis mellifera, colony using DNA microsatellites

Anarchistic behaviour is a very rare phenotype of honeybee colonies. In an anarchistic colony, many workers’ sons are reared in the presence of the queen. Anarchy has previously been described in only two Australian colonies. Here we report on a first detailed genetic analysis of a British anarchistic colony. Male pupae were present in great abundance above the queen excluder, which was clearly indicative of extensive worker reproduction and is the hallmark of anarchy. Seventeen microsatellite loci were used to analyse these male pupae, allowing us to address whether all the males were indeed workers’ sons, and how many worker patrilines and individual workers produced them. In the sample, 95 of 96 of the males were definitely workers’ sons. Given that ≈ 1% of workers’ sons were genetically indistinguishable from queen’s sons, this suggests that workers do not move any queen-laid eggs between the part of the colony where the queen is present to the area above the queen excluder which the queen cannot enter. The colony had 16 patrilines, with an effective number of patrilines of 9.85. The 75 males that could be assigned with certainty to a patriline came from 7 patrilines, with an effective number of 4.21. They were the offspring of at least 19 workers. This is in contrast to the two previously studied Australian naturally occurring anarchist colonies, in which most of the workers’ sons were offspring of one patriline. The high number of patrilines producing males leads to a low mean relatedness between laying workers and males of the colony. We discuss the importance of studying such colonies in the understanding of worker policing and its evolution

Garden varieties: how attractive are recommended garden plants to butterflies?

One way the public can engage in insect conservation is through wildlife gardening, including the growing of insect-friendly flowers as sources of nectar. However, plant varieties differ in the types of insects they attract. To determine which garden plants attracted which butterflies, we counted butterflies nectaring on 11 varieties of summer-flowering garden plants in a rural garden in East Sussex, UK. These plants were all from a list of 100 varieties considered attractive to British butterflies, and included the five varieties specifically listed by the UK charity Butterfly Conservation as best for summer nectar. A total of 2659 flower visits from 14 butterfly and one moth species were observed. We performed a principal components analysis which showed contrasting patterns between the species attracted to Origanum vulgare and Buddleia davidii. The “butterfly bush” Buddleia attracted many nymphalines, such as the peacock, Inachis io, but very few satyrines such as the gatekeeper, Pyronia tithonus, which mostly visited Origanum. Eupatorium cannibinum had the highest Simpson’s Diversity score of 0.75, while Buddleia and Origanum were lower, scoring 0.66 and 0.50 respectively. No one plant was good at attracting all observed butterfly species, as each attracted only a subset of the butterfly community. We conclude that to create a butterfly-friendly garden, a variety of plant species are required as nectar sources for butterflies. Furthermore, garden plant recommendations can probably benefit from being more precise as to the species of butterfly they attract

‘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