Male parentage in dependent-lineage populations of the harvester ant Pogonomyrmex barbatus

We investigated the extent to which workers reproduce in a dependent-lineage population of the monogynous harvester ant Pogonomyrmex barbatus. Dependent-lineage populations contain two interbreeding, yet genetically distinct mitochondrial lineages, each associated with specific alleles at nuclear loci. Workers develop from matings between lineages, and queens develop from matings within lineages, so queens must mate with males of both lineages to produce daughter queens and workers. Males develop from unfertilized eggs and are haploid. Worker production of males could lead to male-mediated gene flow between the lineages if worker-produced males were reproductively capable. This could result in the loss of the dependent-lineage system, because its persistence depends on the maintenance of allelic differences between the lineages. To investigate the extent of worker reproduction in P. barbatus, we genotyped 19-20 males and workers from seven colonies, at seven microsatellite loci, and 1239 additional males at two microsatellite loci. Our methods were powerful enough to detect worker reproduction if workers produced more than 0.39% of males in the population. We detected no worker-produced males; all males appeared to be produced by queens. Thus, worker reproduction is sufficiently infrequent to have little impact on the dependent-lineage system. These results are consistent with predictions based on inclusive fitness theory because the effective queen mating frequency calculated from worker genotypes was 4.26, which is sufficiently high for workers to police those that attempt to reproduce.     

The queen is dead—long live the workers: intraspecific parasitism by workers in the stingless bee Melipona scutellaris

Insect societies are well known for their high degree of cooperation, but their colonies can potentially be exploited by reproductive workers who lay unfertilized, male eggs, rather than work for the good of the colony. Recently, it has also been discovered that workers in bumblebees and Asian honeybees can succeed in entering and parasitizing unrelated colonies to produce their own male offspring. The aim of this study was to investigate whether such intraspecific worker parasitism might also occur in stingless bees, another group of highly social bees. Based on a large-scale genetic study of the species Melipona scutellaris , and the genotyping of nearly 600 males from 45 colonies, we show that ∼20% of all males are workers' sons, but that around 80% of these had genotypes that were incompatible with them being the sons of workers of the resident queen. By tracking colonies over multiple generations, we show that these males were not produced by drifted workers, but rather by workers that were the offspring of a previous, superseded queen. This means that uniquely, workers reproductively parasitize the next-generation workforce. Our results are surprising given that most colonies were sampled many months after the previous queen had died and that workers normally only have a life expectancy of ∼30 days. It also implies that reproductive workers greatly outlive all other workers. We explain our results in the context of kin selection theory, and the fact that it pays workers more from exploiting the colony if costs are carried by less related individuals.     

Lifetime reproductive success and longevity of queens in an annual social insect

Although central to understanding life‐history evolution, the relationship between lifetime reproductive success and longevity remains uncertain in many organisms. In social insects, no studies have reported estimates of queens’ lifetime reproductive success and longevity within populations, despite the importance of understanding how sociality and associated within‐group conflict affect life‐history traits. To address this issue, we studied two samples of colonies of the annual bumblebee, Bombus terrestris audax, reared from wild‐caught queens from a single population. In both samples, queens’ lifetime reproductive success, measured as either queens’ inclusive fitness or as total biomass of queen‐produced sexuals (new queens and males), was significantly positively associated with queen longevity, measured from the day the first worker was produced. We suggest that a positive relationship between reproductive success and longevity was inherited from nonsocial ancestors showing parental care and maintained, at least in part, because the presence of workers buffers queens against extrinsic mortality.     

Optimal timing of the production of sexuals in bumblebee colonies

Colonies of annual eusocial insects often produce only workers until they switch to the production of sexuals (queens and males). This switch from the ergonomic phase to the reproductive phase has been modelled by Macevicz & Oster (1976). Their model predicts that the production of sexuals should take place approximately one generation (one brood developmental period) before resources are depleted in order to maximise the number of sexuals. Bumblebees, however, produce sexuals early (relative to colony lifetime) when resources are still abundant and thus the model of Macevicz & Oster does not hold for these insects.Here we show that Macevicz & Oster's prediction hinges critically on their assumption that the egg laying rate of the queen is a function of the foraging effort of the workers. This assumption does not hold for bumblebees, where queens lay eggs at a constant rate. Taking this feature of bumblebees into account the model predicts early production of sexuals, since it does not pay to produce more workers and collect more resources than needed to raise sexuals.     

Reproductive responses to photoperiod and temperature by artificially hibernated bumblebee (Bombus terrestris) queens

Post‐hibernated bumblebee (Bombus terrestris) queens were kept for 1 week under photoperiodic conditions of 8 h light : 16 h dark, and at four different temperatures (24, 28, 32 and 36°C). The reproductive performance of the queens was then observed. It was found that exposure temperature and hibernation duration did not affect the oviposition rate. The pre‐oviposition period was found to be shortest (3.8 ± 0.7 days) for queens that had hibernated for 4.0 months and had been activated at 28°C. Timing of the initiation of the switch‐point was not affected by exposure temperature and hibernation duration. Significantly higher numbers of workers (268.0 ± 31.4) and sexual queens (119.3 ± 16.8) were produced by the queens that had hibernated for 3.0 months and had been activated at 28 and 36°C, respectively. The queens that had hibernated for 4.0 months and had been activated at 36°C produced the highest number of males (296.2 ± 32.3).     

短舌熊蜂及红光熊蜂耐寒力的测定

以过冷却点和冰点为评价指标,对短舌熊蜂Bombus terrestris的幼虫,蛹,成年工蜂,成年雄蜂,处女蜂王,越冬后的蜂王及红光熊蜂Bombus ignitus的成年工蜂,成年雄蜂,处女蜂王的耐寒力进行检测。结果显示,短舌熊蜂幼虫期耐寒力最低,其次是蛹,工蜂和雄蜂,处女王较高。耐寒力最高的是越冬后的蜂王,其含水量也最低。将短舌熊蜂与红光熊蜂的成年工蜂,成年雄蜂和处女蜂王三型蜂分别进行对比,结果显示,红光熊蜂成年雄蜂与处女蜂王的耐寒力要比短舌熊蜂高。样本的湿重和含水量与过冷却点和冰点无显著相关性。     

Variable worker behaviour in the weakly eusocial sweat bee, <Emphasis Type="Italic">Halictus sexcinctus</Emphasis> Fabricius

Summary 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.     

Effects of photoperiod and hibernation duration on the lifespan of Bombus terrestris

In this study the effects of photoperiod and hibernation duration on the lifespan of Bombus terrestris queens were examined. Hibernation durations of 2.0, 2.5, 3.0, 3.5 and 4.0 months were studied, as were photoperiods of 0 h light : 24 h dark (LD 0:24), LD 8:16, LD 16:8 and LD 24:0. The queens that hibernated for 2.5 months and were exposed to 1 week of LD 8:16 had the highest survival rate (89.3%); the lowest survival rate was found in queens that hibernated for 4.0 months and were reared at LD 24:0. Photoperiod and hibernation duration had significant effects on egg predation by founding queens, competition between queens and workers, and emergence of sexual queens. Hibernation durations of 2.5 and 3.0 months and a photoperiod of LD 8:16 resulted in a significantly longer lifespan of B. terrestris.     

Induction of colony initiation by Japanese native bumble bees using cocoons of the exotic bumblebee Bombus terrestris

The colony initiation rates of Bombus hypocrita (a native Japanese bumblebee) and Bombus terrestris (a European species) foundresses were compared after 4 weeks of exposure to B. terrestris cocoons. The B. terrestris cocoons, when replaced weekly, were effective for inducing oviposition by foundresses of both species. There were no significant differences in the colony initiation rates of B. terrestris and B. hypocrita, either with the control treatment or with the cocoons. The cocoon method was also tested for five species and two subspecies of native Japanese bumblebees. The colony initiation rate was higher for foundresses of the subgenus Bombus s. str. than for foundresses of the subgenera Pyrobombus, Diversobombus, and Thoracobombus. When replaced weekly, the cocoons of B. terrestris are effective inducers of colony foundation in three Japanese native species, namely B. ignitus, B. hypocrita hypocrita, and B. h. sapporoensis.