Young Queens Research Articles

Absence of nepotism in the harassment of duelling queens by honeybee workers.

Nepotism shapes interactions among the members of almost every animal society. However, clear evidence of nepotism within highly cooperative insect societies, such as ant, wasp and honeybee colonies, is rare. Recent empirical findings suggest that nepotism occurs within honeybee colonies where kin-selection theory most strongly predicts its existence: during the lethal queen-queen duels that determine which of several young queens will become the colony's next queen. In this study, I test whether worker bees act nepotistically by hindering duelling queens that are distantly related to themselves. I accomplished this by observing labelled workers harassing duelling queen bees in observation hives and subsequently by determining worker-queen relatedness using DNA microsatellites. I show that the workers that harassed duelling queens were neither more-closely nor more-distantly related to them than were workers selected randomly from the colony. Thus, workers did not behave nepotistically by hindering half-sister queens more than full-sister queens. These results demonstrate that under certain conditions, natural selection limits the evolution of nepotism within animal societies despite strong theoretical predictions for its existence.

Exceptional High Queen Production in the Brazilian Stingless Bee Plebeia remota

In Plebeia remota only a few gynes (young virgin queens) are usually reared per year. However, in two laboratory colonies an extraordinarily high number of gynes emerged within a short period of time. These gynes were significantly smaller than those produced in low numbers by other colonies. Morphometric characters and behaviour of the gynes were recorded. Possible reasons for the phenomenon of exceptional high gyne production are discussed.

Ventilating activity at the hornet nest entrance

The aim of this paper is to provide the first complete description of ventilating activity undertaken by the Oriental hornet. At the portal of each nest of the Oriental hornet (Vespa orientalis, Hymenoptera, Vespinae) there are worker hornets that beat their wings in synchrony with one another, during summer months and at the onset of daylight. These ventilating hornets stand at regular intervals around the nest entrance, not touching, and with their heads facing largely outwards. Although the ventilating hornets stand apart from one another, their wings beat at a uniform pace and there is no connection whatsoever between these hornets and the sentinel worker hornet (or hornets) that inspect those entering the nest. The ventilatory wing beating lasts 3.4 min on average, and upon completion, the ventilating worker either emerges to forage in the field or more often retreats back into the nest. Young queens have seldom been seen ventilating and males have never been seen engaged in such activity. Ordinarily, no ventilatory activity takes place upon the brood combs inside, such as is observed in artificial breeding boxes with a transparent wall, but on rare occasions ventilation may also occur on the inner side of the portal. In cases where there is more than one portal to the nest, the ventilatory activity will occur at each of these sites. Apparently every hornet worker engages in such ventilation at one time or another and their number at any one time is temperature-dependent, that is, the higher the ambient temperature the greater the number the hornets participating in ventilation. Indeed the correlation between the temperature and the ventilatory activity is positive and highly significant.

Tissue localization of the endosymbiotic bacterium "Candidatus Blochmannia floridanus" in adults and larvae of the carpenter ant Camponotus floridanus.

The distribution of endosymbiotic bacteria in different tissues of queens, males, and workers of the carpenter ant Camponotus floridanus was investigated by light and electron microscopy and by in situ hybridization. A large number of bacteria could be detected in bacteriocytes within the midguts of workers, young virgin queens, and males. Large amounts of bacteria were also found in the oocytes of workers and queens. In contrast, bacteria were not present in oocyte-associated cells or in the spermathecae of mature queens, although occasionally a small number of bacteria could be detected in the testis follicles of males. Interestingly, the number of bacteriocytes in mature queens was strongly reduced and the bacteriocytes contained only very few or no bacteria at all, although the endosymbionts were present in huge amounts in the ovaries of the same animals. During embryogenesis of the deposited egg, the bacteria were concentrated in a ring of endodermal tissue destined to become the midgut in later developmental stages. However, during larval development, bacteria could also be detected in other tissues although to a lesser extent. Only in the last-instar larvae were bacteria found exclusively in the midgut tissue within typical bacteriocytes. Tetracycline and rifampin efficiently cleansed C. floridanus workers of their symbionts and the bacteriocytes of these animals still remained empty several months after treatment had ceased. Despite the lack of their endosymbionts, these adult animals were able to survive without any obvious negative effect under normal cultivation conditions.

Production of wax by virgin queens of the stingless bee Melipona bicolor (Apidae, Meliponinae)

Young virgin queens of the stingless bee Melipona bicolor have been shown to produce wax like workers. The small, white to transparent flakes of wax of the queens protrude from the intersegmental space and cover the anterior part of the tergite cuticle, in a way similar to that in workers. This points to the presence of wax glands in the queens. However, workers produce wax from glands located at the fourth to seventh tergites whereas queens secrete wax from the epidermal gland at the third tergite only. Analysis of the queen-produced wax showed that it contains the same substances as the worker-produced wax with minor differences in composition. The wax consists chiefly of the long-chain esters triacontanyl acetate and octacosanyl acetate, smaller amounts of linear C21 to C31 alkanes and alkenes, and still smaller quantities of linear aldehydes and isobutyrate esters. Analysis of wax from wax deposits and wax constructs showed the same composition. Wax from M. bicolor is similar to that of other stingless bees in containing the range of linear long-chain alkenes and alkanes and different from that of Apis bees which contains a more complex mixture, less hydrocarbons and more long chain esters.

Colony development, larval development and worker reproduction in Bombus impatiens Cresson

In this study we describe colony development, larval development, worker reproduction and mating frequency of the queen in laboratory colonies of Bombus impatiens Cresson. Comparison of our observations with data from B. terrestris - the best studied bumblebee - revealed both similarities and dissimilarities between the two. Colonies of B. impatiens have a high number of workers (374.5 ± 108), but relatively low number of young queens (9.4 ± 20) and males (8.9 ± 20). As in B. terrestris, size differences between the castes in B. impatiens were pronounced and became conspicuous as differences in the molting weight of the larvae, starting at the second instar. Workers are able to lay eggs and about 9% of the workers in old colonies had mature oocytes in their ovaries. Despite this, workers were almost never observed to lay eggs in the presence of the queen, and the "competition phase" which is known from B. terrestris was never observed. Mating frequency of the queen was examined using molecular techniques. Two out of ten queens had mated with 2 different males, suggesting some degree of polyandry in the population.

Queen morphology and dispersal tactics in the ant, Cardiocondyla batesii

Dispersal behavior of queens of the myrmicine Cardiocondyla batesii was studied in the springs of 2000 and 2001 in three populations in Southern Spain. Hundreds of young queens were observed to leave their maternal nests, shed their wings, and disperse on foot. Both alate and dealate queens were found to be inseminated, suggesting that they had mated with ergatoid males in their maternal colonies before dispersal. Queens showed a remarkable variation in mesosoma size which appeared to be associated with a polymorphism in wing length (brachyptery vs. macroptery). Morphology did not affect dispersal behavior. Brachyptery is as yet known only from a small number of ant species.

Reduction of Brain Volume Correlates with Behavioral Changes in Queen Ants

The behavior of reproductive female ants distinctly changes during the transition from virgin to mature, egg-laying queen. A winged female ant flies only once during her lifetime when she engages in the nuptial flight. Once she is mated she sheds her wings, excavates a nest and starts laying eggs, the basis for her future colony. We show for two species of harvester ants that this transition is accompanied by changes in the performance of behavioral tests: flying virgins are positively phototactic and prefer open areas, whereas young queens prefer the dark, avoid open areas and, given the opportunity, dig into the soil. These behavioral changes coincide with morphological changes in the brain. The brains of mature queens are significantly smaller than those of virgin females at the time of their mating flight. A disproportionately large shrinkage occurs in the medulla and other parts of the visual system during the early adult life of the queen. The brain reduction appears to be adaptive as mature queens show reduced behavioral repertoires and live in the dark. In contrast to virgin females, they do not rely on vision and might increase their fitness by reducing metabolically costly neural tissue.

The Behavior of Honey Bees (Apis mellifera ligustica) during Queen Duels

Conflict is rare among the members of a highly cooperative society such as a honey bee colony. However, conflict within a colony increases drastically during colony reproduction (‘swarming’) when newly produced queens fight each other until only one queen remains in the nest. This study describes the behavior of queens and workers during naturally occurring queen combat. The duels of five pairs of queens were observed in three observation colonies. A typical duel is described qualitatively and the events of all five duels are described quantitatively. Several aspects of duels that are of particular interest are examined in detail, including the behavior of queens near capped queen cells, worker aggression toward queens, queen tooting, and the relation of queen and worker behavior to the outcome of the duel. The results of this investigation serve as a foundation for rigorous tests of hypotheses regarding the adaptive significance of queen and worker behavior during queen combat. The results presented suggest that: young queens patrol queen cells to kill rival queens while they are vulnerable; workers aggress queens to prevent them from destroying queen cells; queens toot to inhibit worker aggression; workers immobilize queens to make them easy targets for rival queens; and queens eject hind‐gut contents to cause their rival to be immobilized by the workers.

Stenamma debile (Hymenoptera, Formicidae): Productivity and sex allocation across three years

The numerical sex ratios of the facultatively polygynous ant Stenamma debile summed over several collecting sites in southern Germany were 255 males/gyne in 1997, 1.69 males/gyne in 1998 and 5.59 males/gyne in 1999. A large number of female sexuals were produced in queenless colonies, suggesting an inhibitive effect of queens on the production of gynes. Production of males and new workers varied little between the three study years. Production of sexuals plus workers covaried with queen number and worker number. Sex ratio was negatively associated with queen number and positively correlated with total sexual production. Between-year variation of sex ratios is probably correlated with differences in environmental conditions.¶Several mated young queens were found in colonies after the mating seasons in every year. Recruitment of queens was very low in the summer of 1997, and only two of 115 queens from 28 colonies appeared to be young, recently adopted queens. In contrast, 17 young and 59 older queens were collected from 26 colonies in fall 1999. Queen number thus increased by 28.8 % in 1999, but over all three years of 325 queens only 11.4 % were young queens. Incipient colonies with single queens were never observed. Colony structure and productivity of S. debile, as in other ants, apparently vary considerably over periods of several years. Census data from one particular year, and even from three consecutive years, were not sufficient to clarify the causes of variation in sex allocation.

Social parasitism of Psithyrus vestalis in Free-foraging Colonies of Bombus terrestris (Hymenoptera: Apidae)

Females of Psithyrus vestalis (Geoffroy in Fourcroy 1785) were introduced to free-foraging colonies of their host Bombus terrestris (Linnaeus 1758), and the interactions of parasite and host colony members during the development of the colony were studied. In three colonies observed, the B terrestris queens were found dead in their nests between six hours and six days after introduction of the parasite. In two of these colonies the host bees reproduced in males. As all host broods which were reared from eggs which had been laid by workers, only males emerged. In these cases, the Psithyrus females significantly favoured contacts to egg-laying workers, as compared to non-egg-laying workers. The number of contacts to the first group was significantly higher, even before the workers had started laying their eggs. The parasite only reproduced in the third colony, in which the host did not succeed in reproducing young queens or drones.

Sexual reproduction by both queens and workers in the ponerine ant Harpegnathos saltator

Ant workers are very often reproductively degenerate females, but in some species from subfamily Ponerinae, the workers can mate and lay fertilized eggs just like queens (they are then termed "gamergates"). In Harpegnathos saltator, new colonies are founded independently by single queens, and when senescent they are replaced by several gamergates. This secondary polygyny is distinguished by three features: (1) behavioural data indicate that young workers copulate with their brothers, unlike the queens who disperse and mate outside the nests; (2) gamergate colonies produce both male and female sexuals annually; (3) gamergate colonies appear not to fission. Our evidence for the lack of fission is indirect: the nests of H. saltator are unusually elaborate for ponerine ants, and gamergate colonies produce many young queens (at the expense of investing in workers). Queen supersedure results in an extension of colony lifespan, thus making complex constructions possible with a small colony size (65 ± 40 workers in a western Ghats population; N = 59). In turn, these nests represent a valuable resource to be inherited by successive generations of worker offspring. This life history contrasts with that of other ponerine ants having both queens and gamergates (e.g. in Rhytidoponera confusa, nests are simple and short-lived, gamergates are not inbred and their colonies can fission while producing mostly males; Ward, 1981a, 1983a).

The spectacle of intimacy: a public life for the Victorian family

List of Illustrations ix Acknowledgments xi INTRODUCTION The Trouble with Families 3 PART ONE: The Political Theater of Domesticity CHAPTER ONE The Trials of Caroline Norton: Poetry, Publicity, and the Prime Minister 21 CHAPTER TWO: The Young Queen and the Parliamentary Bedchamber: I never saw a man so frightened 46 PART Two: Beneath the Banner of Home t CHAPTER THREE Sarah Stickney Ellis: The Ardent Woman and the Abject Wife 65 CHAPTER FOUR Tom's Pinch: The Sexual Serpent beside the Dickensian Fireside 86 PART THREE: Was That an Angel in the House? CHAPTER FIVE Love after Death: The Deceased Wife's Sister Bill 105 CHAPTER SIX The Transvestite, the Bloomer, and the Nightingale 121 PART FOUR: The Architecture of Comfort and Ruin CHAPTER SEVEN On the Parapets of Privacy: Walls of Wealth and Dispossession 143 CHAPTER EIGHT Robert Kerr: The Gentleman's House and the One-Room Solution 156 PART FIVE: The Sensations of Respectability CHAPTER NINE The Empire of Divorce: Single Women, the Bill of 1857, and Revolt in India CHAPTER TEN Bigamy and Modernity: The Case of Mary Elizabeth Braddon 201 EPILOGUE: Between Manual and Spectacle 215 Notes 221 Index 247

The influence of Nosema apis on young honeybee queens and transmission of the disease from queens to workers

Nosema apis inoculated and non-inoculated honeybee (Apis mellifera) queens were introduced into mating nuclei. The workers accepted all queens from both groups. During the 26 days of the experiment, 47.4% of the inoculated and 50% of the non-inoculated queens were lost. Queens from both groups started egg-laying after 15.8 days. Workers did not perceive the presence of N. apis in their queens and did not supersede inoculated queens significantly more often than healthy ones. Significantly more workers were infected in the mating nuclei with inoculated queens (61%) than in those with non-inoculated ones (5.3%). Apis mellifera / Nosema apis / queen-worker transmission / spore / supersedure

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

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 among queens are predicted to occur. Honey bee colonies reproduce dependently by swarming. The mother queen leaves with a "prime swarm" before daughter queens reach adulthood. Several young queens are produced, and often fight to death after emergence. Surviving queen(s) inherit the established nest or a portion of workers which then depart in an "afterswarm". Honey bee queens show traits considered to be adaptations for fighting and conflict with other queens, such as early venom production and fast development. During fights one of the queens often releases rectal fluid. The function of this "spraying" behaviour is unclear. Possible functions of spraying are to affect worker intervention in fights, to act as a chemical weapon, or to interrupt fights. 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, usually upon physical contact, and it resulted in at least temporary separation of the queens in 81% of the spraying fights. Spraying fights were characterized by a significantly lower proportion of time spent in escalated aggression than non-spraying fights and a significantly shorter first escalated bout. This provides quantitative evidence that spraying interrupts fights and suggests that its function is to provide a temporary respite to the spraying queen.

Does the diapause experience of bumblebee queens Bombus terrestris affect colony characteristics?

Summary1. Bumblebee colonies show much variation in the number of workers, drones, and queens produced. Because this variation prevails even when colonies are kept under identical conditions, it does not seem to be caused by extrinsic factors but rather by differences between founding queens.2. The most likely factor that could cause differences between queens is diapause. Although colonies are raised under standardised conditions, the queens often experience diapause of different length. If there are costs associated with diapause that influence post‐diapause reproduction, the diapause history of the queens could affect colony characteristics.3. Here, several colony characteristics are compared: number of first and second brood workers; total number of workers, drones, and queens; energy spent on sexuals; sex ratio; rate of worker production; time to emergence of first reproductive; and colony lifetime. Colonies were used where the queens experienced a diapause treatment of 0 (nondiapause queens), 2, and 4 months.4. Although no proof was found for the existence of costs associated with diapause, the colony characteristics of nondiapause queens were significantly different from those of diapause queens. Colonies of nondiapause queens produced the lowest number of workers but the highest number of young queens.5. It is argued that these nondiapause colonies are more time‐constrained than diapause colonies because nondiapause colonies produce two generations within the same season and should therefore be more efficient in producing sexual offspring.6. Moreover, nondiapause colonies should rear a more female‐biased sex ratio because they can be certain of the presence of males produced by other (diapause) colonies.

Colony reproduction in Tetragonisca angustula (Apidae, Meliponini)

Colony multiplication of the neotropical stingless bee Tetragonisca angustula started with the finding of a new nesting site by scout bees, followed by the cleaning of the cavity. Cerumen was flown in from the mother colony for the sealing of crevices and the construction of an entrance tube. Major investment of the mother colony in the daughter nest was in workers and cerumen. Almost negligible amounts of pollen were transported from the mother colony after arrival of the gyne in the founding colony. Small quantities of honey were transferred as well. Workers from the daughter nest foraged actively for pollen. Some 500 to 1000 workers entered in the daughter nest, following the virgin queen. This is only a relatively small proportion of the population of the mother colony. Two to four days after the nuptial flight the young queen started egg-laying, and storage pots, mainly filled with honey and some pollen were encountered. In the mother nest, ample pollen and honey stores were present during the months preceding the swarming process. Shortly before swarming took place brood (and adult) populations were 20 to 50% higher than in the other non-swarming colonies, whereas the production of royal cells and males was not different. Data were compared with those got in literature.

Remaking Queen Victoria

List of illustrations Notes on contributors 1. Introduction Margaret Homans and Adrienne Munich Part I. Nation-Making: 2. Nation and nationality: Queen Victoria in the developing narrative of Englishness Elizabeth Langland 3. Crossing the Atlantic with Victoria: American receptions, 1837-1901 Mary Loeffelholz Part II. Queen Victoria and Other Queens: 4. Illustrious company: Victoria among other women in Anglo-American role model anthologies Alison Booth 5. Gloriana Victoriana: Victoria and the cultural memory of Elizabeth I Nicola J. Watson 6. 'Be no more housewives, but queens': Queen Victoria and Ruskin's domestic mythology Sharon Aronofsky Weltman 7. How we lost the empire: retelling the stories of the Rani of Jansi and Queen Victoria Maria Jerinic 8. 'I know what is due to me': self-fashioning and legitimization in Queen Liliuokalani's Hawaii's Story by Hawaii's Queen Robin Bott Part III. Victoria's Career, Early and Late: 9. Reading and writing Victoria: the conduct book and the legal constitution of female sovereignty Gail Turley Houston 10. The wise child and her 'offspring': some changing faces of Queen Victoria Susan P. Casteras '11. I never saw a man so frightened': the young queen and the Parliamentary bedchamber Karen Chase and Michael Levenson 12. The 'Widdy's' empire: Queen Victoria as widow in Kipling's soldier stories and in the Barrack-Room Ballads Dagni Bredeson Part IV. Afterlife: 13. Queen Victoria in the Funnyhouse: Adrienne Kennedy and the rituals of colonial possession Janet Winston Bibliography Index.

Murderous Mothers and The Family/State Analogy in Classical and Renaissance Drama

Sackville and Norton's Gorboduc begins with this summary of its action. The play, written in 1562 during the early years of Elizabeth's reign, is usually interpreted as a piece of advice to the young Queen on matters ranging from the issue of succession to the monarch's dependence on good counsel; having heard of its first performance, Elizabeth commanded a second for her own benefit, clearly recognizing the play's efforts at dutiful counsel. In light of the play's history, then, the portion of the Argument excerpted above is perplexing: rather than attributing the fall of Britain exclusively to Gorboduc's unwise political policies, it immediately shifts focus to the familial relationships at the heart of the kingdom's structure of rule relationships which could, at least apparently, have had little to do with the orphaned Virgin Queen. Gorboduc, the argument clarifies, is not only king and head of the government; he is also a father and husband, and these family ties counterbalance his controlling authority over his kingdom with disastrous results. The country's demise, according to the Argument, is directly traceable to Queen Videna, who more dearly loved her elder son Ferrex, and avenges his death on his brother, thus committing child murder. Rather than foregrounding the king's unwise division of his country which leads to a general uprising

Colony developmental patterns in different local populations of the Turkish bumble bee, Bombus terrestris dalmatinus

SUMMARYAs a preliminary to the use of bumble bee colonies in greenhouse pollination in Turkey, research was initiated on the relevant characteristics of the native bumble bee Bombus terrestris dalmatinus. The same methodology was used that has been successful in the rearing of B. t. terrestris colonies. Queens were collected in four coastal localities (Bodrum, Antalya, Alata and Adana) and characterized by the different dates of the young queen's emergence from diapause. This emergence varied from the end of November to February. Despite the variation in collecting dates, the patterns of development of the colonies produced were similar, indicating that, in this respect, no ecotypes existed. As in B. t. terrestris, the relative timing of the switch point, i.e. when mainly haploid eggs are laid by the queen, is the determining factor for the number of workers and the investment sex ratio of the B. t. dalmatinus colonies. The colonies of these two subspecies develop according to the same principles. The colonies from all four regions had, on average, a male-biased investment sex ratio (0.21, queen fraction), which is a common feature for bumble bees. On the basis of the number of workers produced (c. 150 workers), the colonies from these four Turkish areas are very well suited for pollination purposes.