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In social groups, dominance rank may have important fitness consequences, as higher ranking individuals tend to have higher overall fitness. In social nests of the eastern carpenter bee, Xylocopa virginica, females in social nests demonstrate a complete division of labour where one female is the dominant egg layer and forager while other females in the nest are non-reproductive. We investigated the nature of reproductive queues in this species by performing removal experiments across 3 years to observe how females respond to new reproductive opportunities in the nest. When a primary female was removed, a secondary female always assumed her position as replacement primary and reproductive queues formed in a linear fashion. A third type of female in the nest, the tertiary female, did not become reproductive, even if she was the only female remaining. In delaying reproduction, tertiary females were able to overwinter a second time and were often successful at becoming reproductive in their second summer. Tertiary females were smaller than primary or secondary females, had higher fat stores and lower ovarian development. When all other females in the nest were removed, tertiary females were observed ejecting the offspring of previous dominant females in the nest. Tertiary females appear to represent a novel reproductive strategy among the Hymenoptera who can drastically alter their physiology and behaviour, essentially doubling their life span to maximize reproductive potential.

Female eusocial sweat bees are capable of behaving as queens or workers. Relatively few females become queens, and those that do can directly manipulate the reproductive behaviour of other females in the nest. We collected Lasioglossum (Dialictus) laevissimum workers from nests with and without queens (queenright and queenless nests, respectively) to investigate the influence queens exert on worker behaviour via direct manipulation. Overall, very few L. laevissimum workers (17%) had developed ovaries in Ontario, but queenright and queenless workers were equally likely to have developed ovaries and worn mandibles. However, queenless workers were more likely to be mated than queenright workers. These results suggest first, that queens inhibit egg-laying in most, but not all workers, and second, that queen behaviour during the first few days of workers’ adult lives exerts a lasting influence on worker behaviour. We also compared social traits of L. laevissimum and other Dialictus species using principal components analysis. A strong correlation between worker reproduction and male availability suggests that queen manipulation of the worker brood sex ratio has evolved as an indirect mechanism for queens to discourage worker reproduction.

Sweat bees are one of the most socially polymorphic lineages on the planet. In obligately eusocial species, newly enclosed females may become either queens or workers, depending on the environmental and social circumstances of the nest into which they emerge. In socially polymorphic species, females also have the option of nesting solitarily, founding a nest and raising future reproductives alone, without the help of other adult females. Halictus ligatus is a widespread Nearctic, ground-nesting sweat bee. It has been particularly well studied in Ontario, where detailed studies have described it as obligately eusocial. Here we report evidence that the flexibility of female H. ligatus actually extends to expressing behaviour more typical of socially polymorphic species, those in which some individuals reproduce solitarily. In a population in southern Ontario, black wasps (Astata sp.) emerged from the soil beneath the nesting aggregation and proceeded to excavate their own nesting tunnels, dislocating many H. ligatus nest entrances. Young workers whose natal nests were destroyed by the wasp activity constructed new nests, so under very specific circumstances, it is possible for potential altruists to nest solitarily.

Relocation to novel nests (sometimes called drifting) in flying Hymenoptera is often interpreted as the result of navigation error and guard bees erroneously admitting foreign individuals into the nest. We studied nest fidelity and nest relocation of both females and males in a nesting aggregation of Xylocopa virginica in southern Ontario, Canada, where females can nest either solitarily or socially. Adult female and male bees were trapped at nest entrances, individually paint marked, and then released. Subsequent recapture patterns were used to assess nest fidelity: that is, how faithful individuals were to their home nest and how often they moved to another nest. Bees were considered to have relocated if they were recaptured in a nest different from the one in which they were initially trapped, indicating that they had spent at least one night in a new nest. Some females were only captured in one nest, some occasionally moved to new nests, temporarily or permanently, and a few were never caught in the same nest twice. In addition, females relocated to nests that were further away in 2007 when population density was low, suggesting that they seek out and claim nesting spaces when they are available. Males relocated more frequently than females, with most drifting from nest to nest in no obvious pattern. This indicates that males spend the night wherever space is available or in nests nearest to their territories. This study reveals that for both female and male X. virginica, nest membership is not as stable as once thought.

The bee tribe Ceratinini provides important insights into the early stages of sociality. Many arthropods exhibit behaviours precursory to social life, including adult longevity, parental care, nest loyalty and mutual tolerance, yet the origins of social behaviour are few. Here we describe the biology of a facultatively social bee, Ceratina (Neoceratina) australensis, which exhibits all of the preadaptations for successful group living, yet presents ecological and behavioural characteristics that seemingly disfavour frequent colony formation. This species is socially polymorphic with both solitary and social nests collected in sympatry. Social colonies consist of two foundresses, one contributing both foraging and reproductive effort and the second which remains at the nest as a passive guard. Cooperative nesting provides no overt reproductive benefits over solitary nesting in this population, although brood survival tends to be greater in social colonies. Maternal longevity, subsociality and bivoltine nesting phenology in this species favour colony formation, while dispersal habits and offspring longevity may inhibit more frequent social nesting in this and other ceratinines.

Studies of eusocial halictines suggest that workers have many reproductive options, including sterile altruism in the maternal nest, combined helping and personal reproduction, and diapause and spring nest founding. How and when workers exercise these various options influences the strength of colony social organization. Halictus sexcinctus exhibits highly polymorphic social behaviour, with solitary colonies in central Europe and both eusocial and communal colonies in southern Greece. Indirect evidence suggests that some worker-brood females are actually gynes. A distinctly bimodal size distribution among foundresses in 1998, the lower size peak being close to the modal body size of workers from 1997, suggests that large worker-brood females overwinter and return to the aggregation as eusocial foundresses. Other first-brood females remain in the maternal nest as workers, although few can be classified as classical, sterile altruists. Only 17% of older, healthy workers are sterile (i.e. had ovarian development scores ≤ 0.1), whereas about 83% are reproductive, exhibiting at least one 1/4-developed oocyte. About 57% of older workers have at least one fully or 3/4 developed oocyte, signifying that they are ready or almost ready to lay. Sterile workers exhibit greater total wear (combined mandibular and wing wear) scores than reproductive workers, suggesting that they are older, have higher activity rates, or both.

We studied the nesting and social biology of two aggregations of the obligately eusocial halictine bee Lasioglossum malachurum at Agios Nikolaos Monemvasias (ANM) in southern Greece. Observations and nest excavations carried out from May to June 2000, revealed social and demographic variation between aggregations and years at ANM, as well as notable differences between these and other European populations. In southern Greece, the colony cycle includes multiple broods: the first two broods comprise only workers, whereas the third brood comprises workers, males, and gynes. Although pleometrosis is unknown in other populations, in the ANM region, as many as 10% of nests have more than one foundress. Newly emerged workers and gynes exhibit non-overlapping size distributions, but a few queens are worker-sized, indicating that workers occasionally overwinter and become foundresses. Although the vast majority of workers are unmated and most exhibit no ovarian development, an increase in worker ovarian development at the time of male production suggests that many males may develop from worker-laid eggs. Worker reproduction seems to be inhibited by the presence of queens, and annual variation in queen mortality may underlie annual variation in worker ovarian development. Across Europe, the major demographic and social differences among L. malachurum populations are in the number of worker broods and the extent of worker ovarian development. This contrasts with the results of a principal components analysis of social traits among 15 social L. (Evylaeus) populations, which shows that interspecific social variation is defined by the proportion of males in the early brood, the proportion of workers mated, queen-worker size dimorphism, gyne overwintering locale, and the proportion of workers with developed ovaries.

Female sweat bees in the species Halictus ligatus exhibit a wide range of reproductive roles, ranging from typically foundress or queen-like to typically worker-like. Nests are founded in spring and most are haplometrotic, that is, founded by a single foundress. A few (up to 12 %) are pleometrotic, founded by 2-6 foundresses. Variation in the proportion of multifoundress nests from year to year and from place to place suggests an adaptive basis for pleometrosis. We studied the demographic and social characteristics of 23 pleometrotic nests in an aggregation of 250-300 nests near Victoria, Ontario, in 1984, 1990, and 1991. In pleometrotic associations, dominant foundresses behaved in a manner typical of mid-summer, haplometrotic queens, while subordinates behaved like mid-summer workers. Dominant foundresses tended to be larger than subordinates. Pleometrotic nests were significantly more likely than haplometrotic nests to produce brood, and they also produced more workers. However, this early advantage did not result in the production of more reproductive brood per nest, nor did pleometrotic foundresses experience higher productivity per foundress than did haplometrotic foundresses. Relatively low relatedness among various categories of brood implied that subordinate foundresses were not closely related to dominants. We suggest that pleometrosis most likely results from accidental encounters between spring foundresses as they leave their hibernacula. Once formed, such associations confer a survival advantage on the nest as a whole, but do not result in greater reproductive brood productivity.

Halictine bees exhibit an enormous diversity of solitary and social colony structures. To investigate social evolution in the genusHalictus, phylogenies of 15 species of the subgeneraH. (Halictus) andH. (Seladonia) were constructed based on protein electrophoretic data. Solitary, social, and socially polymorphic species were included.Halictus (Seladonia) apparently rendersH. (Halictus) paraphyletic. The common ancestor ofH. (Halictus) andH. (Seladonia) was probably social or socially polymorphic. This implies that some solitary and socially polymorphic species, such asH. confusus andH. tumulorum, represent evolutionary reversals from a completely eusocial condition to the solitary condition that is thought to be primitive for the subfamily as a whole.