Queens In Colonies Research Articles

Queen execution increases relatedness among workers of the invasive Argentine ant, Linepithema humile.

Polygyny in social insects can greatly reduce within-nest genetic relatedness. In polygynous ant species, potential rival queens in colonies with multiple queens are often executed by other queens, workers, or both. The Argentine ant, Linepithema humile, native to South America, forms a "supercolony" that is composed of a large number of nests and is considered to contribute to the ant's invasion success. Currently, four mutually antagonistic supercolonies are contiguously distributed within a small area of Japan. Here, we analyzed the genetic structure and relatedness within and among the four supercolonies using microsatellite markers to clarify how L.humile maintains its supercoloniality. The results of AMOVA and BASP, the F ST values, and the existence of several private alleles indicated that the L.humile population in the Kobe area had a characteristic genetic structure. Within a given supercolony, there was significant genetic differentiation (F ST) among workers collected in May and those collected in September. The significant deviation from Hardy-Weinberg equilibrium increased, and the relatedness among workers significantly increased from May to September in all supercolonies. This result suggested that the supercolonies replaced old queens with new ones during the reproductive season, thus supporting the plausibility of queen execution. From the perspective of kin selection, workers collectively eliminate queens, thereby increasing their own inclusive fitness. Restricted gene flow among supercolonies, together with mating with sib and queen execution, could help to maintain the unique social structure of L.humile, the distribution of which is expanding worldwide.

Octopamine and cooperation: octopamine regulates the disappearance of cooperative behaviours between genetically unrelated founding queens in the ant.

We investigated whether octopamine (OA) is associated with the disappearance of cooperation in Polyrhachis moesta ant queens. Queens of P. moesta facultatively found the colony with genetically unrelated queens. The founding queens perform frequent food exchange with these non-related queens and partake in cooperative brood rearing, whereas single colony queens exclude non-related queens via aggressive behaviour. Thus, aggression is a factor that reduces cooperation. Given that aggression is generally associated with brain OA in insects, we hypothesized that OA controls the behavioural change in cooperation in the ant queen, via an increase in aggression. To test this hypothesis, we compared the amounts of OA and related substances in the brain between founding and colony queens, and observed the interaction of founding queens following oral OA administration. The brain OA levels in colony queens were significantly higher than those in founding queens. Oral administration of OA to founding queens caused significantly less trophallaxis and allogrooming behaviour than in the control founding queens, but with no significant increase in aggression. These results suggest that OA promotes the disappearance of cooperation in founding queens of P. moesta. This is the first study to reveal the neuroendocrine mechanism of cooperation in ant queens.

Multiple functions of fire ant Solenopsis invicta mandibular gland products

AbstractAlarm pheromones of social insects are best known for their role in the defence and maintenance of colony integrity. Previous studies with the fire ant Solenopsis invicta Buren (Hymenoptera: Formicidae) demonstrate that the mandibular glands of workers (sterile females) and male and female sexuals produce an alarm pheromone, 2‐ethyl‐3,6‐dimethylpyrazine. The function of alarm pheromones in worker ants is well understood and divergent from the function of these compounds in the winged sexual forms. The present study quantifies the amount of pyrazine in the mandibular glands from male and female alate sexuals, as well as queens. Pyrazine production in female alates starts in the late pupal stage and increases until they reach mating flight‐ready maturity; however, after mating flight participation, the pyrazine level declines by >50%. Interestingly, mature male alates lose >85% of their mandibular gland pyrazine during mating flight activity. The results of the present study indicate that male and female sexuals use mandibular gland secretions for mating flight initiation and during mating flights. Furthermore, the ontogeny of mandibular gland products (pyrazine as the marker) from newly‐mated queens to mature colony queens shows a more than two‐fold increase in the amount of pyrazine by 6 months after mating. However, this is followed by a decline to trace amounts in mature colony queens (>2 years old), suggesting a function for mandibular gland products during colony development. Multifunctional use of social insect pheromones is well documented and data are reported in the present study suggesting new roles for mandibular gland products in fire ants.

Mating frequencies of honey bee queens (Apis mellifera L.) in a population of feral colonies in the Northeastern United States.

Across their introduced range in North America, populations of feral honey bee (Apis mellifera L.) colonies have supposedly declined in recent decades as a result of exotic parasites, most notably the ectoparasitic mite Varroa destructor. Nonetheless, recent studies have documented several wild populations of colonies that have persisted. The extreme polyandry of honey bee queens—and the increased intracolony genetic diversity it confers—has been attributed, in part, to improved disease resistance and may be a factor in the survival of these populations of feral colonies. We estimated the mating frequencies of queens in feral colonies in the Arnot Forest in New York State to determine if the level of polyandry of these queens is especially high and so might contribute to their survival success. We genotyped the worker offspring from 10 feral colonies in the Arnot Forest of upstate New York, as well as those from 20 managed colonies closest to this forest. We found no significant differences in mean mating frequency between the feral and managed queens, suggesting that queens in the remote, low-density population of colonies in the Arnot Forest are neither mate-limited nor adapted to mate at an especially high frequency. These findings support the hypothesis that the hyperpolyandry of honey bees has been shaped on an evolutionary timescale rather than on an ecological one.

Social structure varies with elevation in an Alpine ant.

Insect societies vary greatly in social organization, yet the relative roles of ecological and genetic factors in driving this variation remain poorly understood. Identifying how social structure varies along environmental gradients can provide insights into the ecological conditions favouring alternative social organizations. Here, we investigate how queen number variation is distributed along elevation gradients within a socially polymorphic ant, the Alpine silver ant Formica selysi. We sampled low- and high-elevation populations in multiple Alpine valleys. We show that populations belonging to different drainage basins are genetically differentiated. In contrast, there is little genetic divergence between low- and high-elevation populations within the same drainage basin. Thus, elevation gradients in each of the drainage basins represent independent contrasts. Whatever the elevation, all well-sampled populations are socially polymorphic, containing both monogynous (= one queen) and polygynous (=multiple queen) colonies. However, the proportion of monogynous colonies per population increases at higher elevation, while the effective number of queens in polygynous colonies decreases, and this pattern is replicated in each drainage basin. The increased prevalence of colonies with a single queen at high elevation is correlated with summer and winter average temperature, but not with precipitation. The colder, unpredictable and patchy environment encountered at higher elevations may favour larger queens with the ability to disperse and establish incipient monogynous colonies independently, while the stable and continuous habitat in the lowlands may favour large, fast-growing polygynous colonies. By highlighting differences in the environmental conditions favouring monogynous or polygynous colonies, this study sheds light on the ecological factors influencing the distribution and maintenance of social polymorphism.

Size variation in eggs laid by normal-sized and miniature queens of Plebeia remota (Holmberg) (Hymenoptera: Apidae: Meliponini)

Miniature stingless bee queens have been studied concerning frequency distribution, production and egg laying performance. This study aimed to investigate size variation in eggs laid by Plebeia remota (Holmberg) queens and whether it is due to differences in queen size or colony conditions. A sample of 10 queens (8 of typical size and 2 miniature) was measured morphometrically (head width, interorbital distance, and intertegular distance) as well the eggs they laid (length, width and volume). Initially, eggs were analyzed when laid by queens in their own colonies. Significant differences were found for length, width and volume of eggs considering the total group of queens or both queen morphotypes. However, no significant correlations were found between queen size and egg size. Afterwards, two experiments were performed to evaluate the influence of colony conditions on egg size. Firstly, we shifted the queens from their original colonies (i.e., a typical queen was placed into a miniature queen colony, and vice-versa). Secondly, they were put into another colony (both types of queens, one each time, were placed on a third colony, a ‘host colony’). In all situations, both queen morphotypes laid eggs of similar or different sizes than before, often with significant differences. The results indicate that variation in egg size is due to conditions imposed to queens in the colony (e.g. queen feeding status, number of cells available to be oviposited), and not due to variation in queen body size.

Convergent Evolution: The Genetics of Queen Number in Ants

Large, non-recombining genomic regions underlie the polymorphism in colony queen number in two distantly related ant species. This illustrates that convergence in complex phenotypes can arise via convergence in general genomic architecture, rather than convergent changes in specific genes.

The Effect of Open Brood and Colony Strength on the Onset of Oviposition by Queen Bees

Abstract In bee colonies without open brood, e.g., after swarming, there is no need for royal jelly, and nurse bees thus do not produce it. According to many beekeepers, adding combs with open brood restarts the production of royal jelly by nurse bees, and the virgin queens then are better fed and start earlier oviposition. The purpose of this study was to investigate whether the presence of open brood and the strength of the colonies affect the onset of oviposition by queen bees. Open brood in colonies with virgins before and during mating flights did not accelerate the initiation of oviposition by the queens. In addition, no differences were identified in starting oviposition by queens in strong colonies of more than 30,000 worker bees, or in weak colonies with up to 1,000 workers. Overall, the results showed that neither open brood in the nests, nor the strength of the colonies affects the onset of oviposition by queen bees.

Strength in numbers: Large and permanent colonies have higher queen oviposition rates in the invasive Argentine ant (Linepithema humile, Mayr)

Polydomy associated with unicoloniality is a common trait of invasive species. In the invasive Argentine ant, colonies are seasonally polydomous. Most follow a seasonal fission–fussion pattern: they disperse in the spring and summer and aggregate in the fall and winter. However, a small proportion of colonies do not migrate; instead, they inhabit permanent nesting sites. These colonies are large and highly polydomous. The aim of this study was to (1) search for differences in the fecundity of queens between mother colonies (large and permanent) and satellite colonies (small and temporal), (2) determine if queens in mother and satellite colonies have different diets to clarify if colony size influences social organization and queen feeding, and (3) examine if colony location relative to the invasion front results in differences in the queen’s diet. Our results indicate that queens from mother nests are more fertile than queens from satellite nests and that colony location does not affect queen oviposition rate. Ovarian dissections suggest that differences in ovarian morphology are not responsible for the higher queen oviposition rate in mother vs. satellite nests, since there were no differences in the number and length of ovarioles in queens from the two types of colonies. In contrast, the higher δ15N values of queens from mother nests imply that greater carnivorous source intake accounts for the higher oviposition rates.

Orphaning does not affect the colony productivity of the primitive eusocial wasp Polistes snelleni

In colonies of primitively eusocial wasps, some dominant workers become successive queens and inherit queenship after the death of the foundress queens. Although workers in many species do not mate, workers of Polistes snelleni are capable of mating and female production. In this study, we removed foundress queens from colonies of P. snelleni to evaluate the effects of queen loss on the dominant–subordinate relationships among the remaining workers and the productivity of colonies in the species. The foundress queens were the sole egg layers in almost all of the queenright colonies. The frequency of dominance behaviour among the wasps in the queenright colonies was significantly less than in the orphan colonies. The frequency of dominance behaviour in the successive queens after queen removal was significantly more than in the foundress queens. Multiple workers had developed ovaries, including the successive queens in 66.7 % (10/15) of the orphan colonies after queen removal. The orphan colonies produced significantly more cells and eggs than the queenright colonies. Our results suggest that the reproductive potential of the successive queens in the orphan colonies is not lower than that of the foundress queens, and that the productivity of the orphan colonies is maintained rather than causing potential conflict over direct reproduction among workers.

Shared genes related to aggression, rather than chemical communication, are associated with reproductive dominance in paper wasps (Polistes metricus)

BackgroundIn social groups, dominant individuals may socially inhibit reproduction of subordinates using aggressive interactions or, in the case of highly eusocial insects, pheromonal communication. It has been hypothesized these two modes of reproductive inhibition utilize conserved pathways. Here, we use a comparative framework to investigate the chemical and genomic underpinnings of reproductive dominance in the primitively eusocial wasp Polistes metricus. Our goals were to first characterize transcriptomic and chemical correlates of reproductive dominance and second, to test whether dominance-associated mechanisms in paper wasps overlapped with aggression or pheromone-related gene expression patterns in other species. To explore whether conserved molecular pathways relate to dominance, we compared wasp transcriptomic data to previous studies of gene expression associated with pheromonal communication and queen-worker differences in honey bees, and aggressive behavior in bees, Drosophila, and mice.ResultsBy examining dominant and subordinate females from queen and worker castes in early and late season colonies, we found that cuticular hydrocarbon profiles and genome-wide patterns of brain gene expression were primarily associated with season/social environment rather than dominance status. In contrast, gene expression patterns in the ovaries were associated primarily with caste and ovary activation. Comparative analyses suggest genes identified as differentially expressed in wasp brains are not related to queen pheromonal communication or caste in bees, but were significantly more likely to be associated with aggression in other insects (bees, flies), and even a mammal (mice).ConclusionsThis study provides the first comprehensive chemical and molecular analysis of reproductive dominance in paper wasps. We found little evidence for a chemical basis for reproductive dominance in P. metricus, and our transcriptomic analyses suggest that different pathways regulate dominance in paper wasps and pheromone response in bees. Furthermore, there was a substantial impact of season/social environment on gene expression patterns, indicating the important role of external cues in shaping the molecular processes regulating behavior. Interestingly, genes associated with dominance in wasps were also associated with aggressive behavior in bees, solitary insects and mammals. Thus, genes involved in social regulation of reproduction in Polistes may have conserved functions associated with aggression in insects and other taxa.

The effect of one generation of controlled mating on the expression of hygienic behaviour in honey bees

SummaryHoney bee mating cannot be directly controlled in the same way as in many agriculturally important animals. Instrumental insemination is, however, possible and can be used as an aid in selective breeding. Hygienic behaviour, in which worker bees detect and remove dead or diseased brood from capped cells, is a heritable trait that confers colony-level resistance against brood diseases. Using the freeze-killed brood (FKB) bioassay we compared the levels of hygiene in colonies headed by daughter queens reared from hygienic mother colonies that were either instrumentally inseminated with sperm from drones reared from hygienic colonies or allowed to mate naturally with naturally-occurring drones. Hygiene levels were significantly higher in the colonies of the instrumentally inseminated queens than in the colonies of the naturally-mated queens. However, the hygiene levels in the naturally-mated colonies were encouragingly high and indicate that supplying beekeepers with naturally-mated queens, or virgin queens to mate locally, can result in colonies with high levels of hygiene.

A test of neuroecological predictions using paperwasp caste differences in brain structure (Hymenoptera: Vespidae)

Adaptive brain architecture hypotheses predict brain region investment matches the cognitive and sensory demands an individual confronts. Social hymenopteran queen and worker castes differ categorically in behavior and physiology leading to divergent sensory experiences. Queens in mature colonies are largely nest-bound while workers depart nests to forage. We predicted social paperwasp castes would differ in tissue allocation among brain regions. We expected workers to invest relatively more than queens in neural tissues that process visual input. As predicted, we found workers invested more in visual relative to antennal processing than queens both in peripheral sensory lobes and in central processing brain regions (mushroom bodies). Although we did not measure individual brain development changes, our comparative data provide a preliminary test of mechanisms of caste differences. Paperwasp species differ in the degree of caste differentiation (monomorphic versus polymorphic castes) and in colony structure (independent- versus swarm-founding); these differences could correspond to the magnitude of caste brain divergence. If caste differences resulted from divergent developmental programs (experience-expectant brain growth), we predicted species with morphologically distinct queens, and/or swarm-founders, would show greater caste divergence of brain architecture. Alternatively, if adult experience affected brain plasticity (experience-dependent brain growth), we predicted independent-founding species would show greater caste divergence of brain architecture. Caste polymorphism was not related to the magnitude of queen-worker brain differences, and independent-founder caste brain differences were greater than swarm-founder caste differences. Greater caste separation in independent-founder brain structure suggests a role for adult experience in the development of caste-specific brain anatomy.

The value of oviposition timing, queen presence and kinship in a social insect

Reproductive cooperation confers benefits, but simultaneously creates conflicts among cooperators. Queens in multi-queen colonies of ants share a nest and its resources, but reproductive competition among queens often results in unequal reproduction. Two mutually non-exclusive factors may produce such inequality in reproduction: worker intervention or queen traits. Workers may intervene by favouring some queens over others, owing to either kinship or queen signals. Queens may differ in their intrinsic fecundity at the onset of oviposition or in their timing of the onset of oviposition, leading to their unequal representation in the brood. Here, we test the role of queen kin value (relatedness) to workers, timing of the onset of oviposition and signals of presence by queens in determining the maternity of offspring. We show that queens of the ant Formica fusca gained a significantly higher proportion of sexuals in the brood when ovipositing early, and that the presence of a caged queen resulted in a significant increase in both her share of sexual brood and her overall reproductive share. Moreover, the lower the kin value of the queen, the more the workers invested in their own reproduction by producing males. Our results show that both kinship and breeding phenology influence the outcome of reproductive conflicts, and the balance of direct and indirect fitness benefits in the multi-queen colonies of F. fusca.

Altruistic Behavior by Egg-Laying Worker Honeybees

If a honeybee (Apis mellifera) colony loses its queen, worker bees develop their ovaries and produce male offspring [1]. Kin selection theory predicts that the degree of altruism in queenless colonies should be reduced because the relatedness of workers to a hivemate's offspring is less in queenless colonies than it is to the daughters of the queen in queenright colonies [2-4]. To explore this hypothesis, we examined the behavior and physiology of queenless egg-laying workers. Queenless bees engaged in both personal reproduction and the social foraging and defense tasks that benefited their colony. Laying workers also had larger brood-food-producing and wax glands, showing metabolic investments in both colony maintenance and personal reproduction. Whereas in queenright colonies there is a very clear age-based pattern of division of labor between workers, in queenless colonies the degree of individual specialization was much reduced. Queenless colonies functioned as a collective of reproductive and behaviorally generalist bees that cooperatively maintained and defended their nest. This social structure is similar to that observed in a number of primitively social bee species [5]. Laying workers therefore show a mix of selfish personal reproduction and altruistic cooperative behavior, and the queenless state reveals previously unrecognized plasticity in honeybee social organization.

Regional variation in queen and worker aggression in incipient colonies of the desert ant Messor pergandei

The fate of queen foundress associations in ants varies across taxa: in some, lethal fighting results in survival of a single queen, while in others, queens coexist long term. One hypothesis for this difference is that selection favors fighting when group sizes are small and tolerance when groups are large. In an experiment with the ant Messor pergandei, we formed small, medium, and large groups with newly mated queens from field populations that have different mean group sizes and differ in whether multiple queens occur in older established colonies. We found that whether queens are eliminated by fighting depends upon region of origin and not group size: regardless of co-foundress number, queens from sites with single-queen adult field colonies displayed agonistic behaviors and their colonies reduced to a single queen, while queens from sites with multiple-queen colonies did not fight and co-foundresses coexisted long term. Worker aggression towards and elimination of queens were also correlated with region of origin. Where fighting occurred, queens were as likely to be killed by workers as by other queens. An aggressive display was the most common form of agonistic interaction among queens, while fighting was relatively rare. We hypothesize that queen displays evolved in response to worker attacks because they increase the probability that workers will eliminate competitor queens. Our results suggest that the evolutionary interests of workers, as well as queens, could be important in determining the evolution and maintenance of queen elimination in foundress associations.

Chemical identity of recently emerged workers, males, and queens in the stingless bee Melipona marginata

Epicuticular hydrocarbons are involved in several behavioral processes in social insects including recognition of nestmates, castes, and sex. The aim of the present study was to characterize and to verify differences in the chemical profiles of 1-day recently emerged workers, males, and queens in colonies of the stingless bee Melipona marginata. Gas chromatography–mass spectrometry and multivariate analyses of bee samples from three colonies clearly separated cuticular hydrocarbon profiles of males, queens, and workers. The results showed that cuticular hydrocarbon profiles were comprised of alkanes, alkenes, and methyl alkanes that varied quantitatively and qualitatively according to the castes and sex. Different from previous studies, our investigation also demonstrated that males and queens presented higher similarity in their epicuticular hydrocarbons than to workers.

The long-lived queen: reproduction and longevity in female eusocial Damaraland mole-rats (Fukomys damarensis)

The inverse relationship between reproduction and lifespan is one of the main concepts of life history theory. This association has been observed in most taxa, although exceptions have been found in which a breeding female outlives her non-reproductive cohorts. This relationship is well known in social insects, and it has recently come to light that reproductive females of certain social mole-rat species also exhibit extended lifespans relative to non-breeders. We analysed mark-recapture data over 13 years in 12 colonies of Damaraland mole-rats (Fukomys damarensis) to assess if colony queens lived longer than non-reproductive adult females. Queens were recaptured up to 8.5 years after initial capture (X = 6.2 years); significantly longer than non-reproductive females (X = 1.3 years), suggesting that a colony queen lives longer than her non-reproductive subordinates. This difference may be attributed to both physiological and social factors which may relax reproductive costs in queens.

The Long-Lived Queen: Reproduction and Longevity in Female Eusocial Damaraland Mole-Rats (Fukomys damarensis)

The inverse relationship between reproduction and lifespan is one of the main concepts of life history theory. This association has been observed in most taxa, although exceptions have been found in which a breeding female outlives her non-reproductive cohorts. This relationship is well known in social insects, and it has recently come to light that reproductive females of certain social mole-rat species also exhibit extended lifespans relative to non-breeders. We analysed mark-recapture data over 13 years in 12 colonies of Damaraland mole-rats (Fukomys damarensis) to assess if colony queens lived longer than non-reproductive adult females. Queens were recaptured up to 8.5 years after initial capture (X = 6.2 years); significantly longer than non-reproductive females (X = 1.3 years), suggesting that a colony queen lives longer than her non-reproductive subordinates. This difference may be attributed to both physiological and social factors which may relax reproductive costs in queens.

Polygyny in the nest-site limited acacia-ant Crematogaster mimosae

Polygyny is common in social insects despite inevitable decreases in nestmate relatedness and reductions to the inclusive fitness returns for cooperating non-reproductive individuals. We studied the prevalence and mode of polygyny in the African acacia-ant Crematogaster mimosae. These ants compete intensively with neighboring colonies of conspecifics and with three sympatric ant species for resources associated with the whistling-thorn acacias in which they all obligately nest. We used the genotypes of alate males at ten microsatellite loci to reconstruct queen genotypes and found that C. mimosae colonies are frequently secondarily polygynous, in that they include multiple closely related (and sometimes full-sib) queens, and (more rarely) unrelated queens. We also found that individual queens in both monogynous and polygynous colonies had mated with multiple males, making C. mimosae an interesting example of simultaneous polygyny and polyandry. The presence of polygyny in C. mimosae and the intense competition for nest-sites between C. mimosae and its conspecifics support the association between nest-site limitation and polygyny. Polygyny may allow for increased worker populations and a competitive advantage, as inter-colony conflicts are typically won by the colony with the larger number of workers.