Rosalyn S. Gloag

A molecular phylogeny of the genus Apis suggests that the Giant Honey Bee of the Philippines, A. breviligula Maa, and the Plains Honey Bee of southern India, A. indica Fabricius, are valid species

Two new taxa have been added to the existing molecular phylogenies of the genus Apis. The new phylogeny supports the accepted phylogenetic relationships of {dwarf honey bees [giant honey bees (cavity‐nesting honey bees)]}. Based on Bayesian and maximum parsimony trees, our analysis supports recognition of Apis indica, the Plains Honey Bee of south India, as a separate species from A. cerana. Our analysis also supports recognition of the Giant Philippines Honey Bee, A. breviligula, as a separate species from A. dorsata.

Social parasitism by workers in queenless and queenright Apis cerana colonies.

We examined worker reproduction in queenless and queenright Apis cerana colonies to determine if they are parasitized by workers from other nests. The results demonstrate that 2–6% of workers in queenright colonies are from another nest (non‐natal), but these workers are not statistically more likely to have activated ovaries than natal workers, and are therefore unlikely to be active parasites. However, in queenless colonies we found a significant difference between the proportion of non‐natal (72.7%) and natal (36.3%) workers with activated ovaries. Non‐natal workers also had significantly higher reproductive success than natal workers: 1.8% of workers were non‐natal, but these laid 5.2% of the eggs and produced 5.5% of the pupae. Unlike A. florea, the proportion of non‐natal workers does not increase in queenless nests.

Thelytokous Parthenogenesis in Unmated Queen Honeybees (Apis mellifera capensis): Central Fusion and High Recombination Rates

The subspecies of honeybee indigenous to the Cape region of South Africa, Apis mellifera capensis, is unique because a high proportion of unmated workers can lay eggs that develop into females via thelytokous parthenogenesis involving central fusion of meiotic products. This ability allows pseudoclonal lineages of workers to establish, which are presently widespread as reproductive parasites within the honeybee populations of South Africa. Successful long-term propagation of a parthenogen requires the maintenance of heterozygosity at the sex locus, which in honeybees must be heterozygous for the expression of female traits. Thus, in successful lineages of parasitic workers, recombination events are reduced by an order of magnitude relative to meiosis in queens of other honeybee subspecies. Here we show that in unmated A. m. capensis queens treated to induce oviposition, no such reduction in recombination occurs, indicating that thelytoky and reduced recombination are not controlled by the same gene. Our virgin queens were able to lay both arrhenotokous male-producing haploid eggs and thelytokous female-producing diploid eggs at the same time, with evidence that they have some voluntary control over which kind of egg was laid. If so, they are able to influence the kind of second-division meiosis that occurs in their eggs post partum.

Nest site selection in the open-nesting honeybee Apis florea

We studied nest site selection by swarms of the red dwarf honeybee, Apis florea. By video recording and decoding all dances of four swarms, we were able to determine the direction and distances indicated by 1,239 dances performed by the bees. The bees also performed a total of 715 nondirectional dances; dances that were so brief that no directional information could be extracted. Even though dances converged over time to a smaller number of areas, in none of the swarms did dances converge to one site. As a result, even prior to lift off, bees performed dances indicating nest sites in several different directions. Two of four swarms traveled directly in what seemed to be the general direction indicated by the majority of dances in the half hour prior to swarm lift off. The other two traveled along circuitous routes in the general direction indicated by the dances. We suggest that nest site selection in A. florea has similar elements to nest site selection in the better-studied Apis mellifera. However, the observation that many more locations are indicated by dances prior to lift off also shows that there are fundamental differences between the two species.

Maternity of replacement queens in the thelytokous Cape honey bee Apis mellifera capensis

Unlike workers of all other honey bee (Apis mellifera) subspecies, workers of the Cape honey bee of South Africa (A. mellifera capensis) reproduce thelytokously and are thus able to produce female offspring that are pseudoclones of themselves. This ability allows workers to compete with their queen over the maternity of daughter queens and, in one extreme case, has led to a clonal lineage of workers becoming a social parasite in commercially managed populations of A. mellifera scutellata. Previous work (Jordan et al., Proc R Soc Lond B Biol Sci 275:345, 2008) showed that, in A. mellifera capensis, 59% of queen cells produced during swarming events contained the offspring of workers and that, of these, 65% were the offspring of non-natal workers. Here, we confirm that a substantial proportion (38.5%) of offspring queens is worker-laid. We additionally show that: (1) Although queens produce most diploid female offspring sexually, we found some homozygous or hemizygous queen offspring, suggesting that queens also reproduce by thelytoky. These parthenogenetic individuals are probably nonviable beyond the larval stage. (2) Worker-laid offspring queens are viable and become the resident queen at the same frequency as do sexually produced queen-laid offspring queens. (3) In this study, all but one of the worker-derived queens were laid by natal workers rather than workers from another nest. This suggests that the very high rates of social parasitism observed in our previous study were enhanced by beekeeping manipulations, which increased movement of parasites between colonies.

Dance precision of Apis florea—clues to the evolution of the honeybee dance language?

All honeybee species make use of the waggle dance to communicate the direction and distance to both food sources and potential new nest sites. When foraging, all species face an identical problem: conveying information about profitable floral patches. However, profound differences in nesting biology (some nest in cavities while others nest in the open, often on a branch or a cliff face) may mean that species have different requirements when dancing to advertise new nest sites. In cavity nesting species, nest sites are a precise location in the landscape: usually a small opening leading to a cavity in a hollow tree. Dances for cavities therefore need to be as precise as possible. In contrast, when the potential nest site comprises a tree or perhaps seven a patch of trees, precision is less necessary. Similarly, when a food patch is advertised, dances need not be very precise, as floral patches are often large, unless they are so far away that recruits need more precise information to be able to locate them. In this paper, we study the dance precision of the open-nesting red dwarf bee Apis florea. By comparing the precision of dances for food sources and nest sites, we show that A. florea workers dance with the same imprecision irrespective of context. This is in sharp contrast with the cavity-nesting Apis mellifera that increases the precision of its dance when advertising a potential new home. We suggest that our results are in accordance with the hypothesis that the honeybees’ dance communication initially evolved to convey information about new nest sites and was only later adapted for the context of foraging.

Nest defence in a stingless bee: What causes fighting swarms in Trigona carbonaria (Hymenoptera, Meliponini)?

Abstract.The Australian stingless bee Trigona carbonaria sometimes displays a striking collective behaviour, known as a ‘fighting swarm’ in which thousands of workers fight and die. Molecular analysis of eight naturally-occurring fights showed they almost always comprise just two colonies, one of which is located within 2 m of the fight. Fighting swarms were experimentally triggered by manipulating colonies so that they received non-nestmate workers. Combined, our investigations suggest that T. carbonaria fighting swarms arise as a collective defence of the nest from conspecific invasion (e.g. robbery or nest usurpation)

An invasive social insect overcomes genetic load at the sex locus

Some invasive hymenopteran social insects found new populations with very few reproductive individuals. This is despite the high cost of founder effects for such insects, which generally require heterozygosity at a single locus—the complementary sex determiner, csd—to develop as females. Individuals that are homozygous at csd develop as either infertile or subfertile diploid males or not at all. Furthermore, diploid males replace the female workers that are essential for colony function. Here we document how the Asian honey bee (Apis cerana) overcame the diploid male problem during its invasion of Australia. Natural selection prevented the loss of rare csd alleles due to genetic drift and corrected the skew in allele frequencies caused by founder effects to restore high average heterozygosity. Thus, balancing selection can alleviate the genetic load at csd imposed by severe bottlenecks, and so facilitate invasiveness.

No worker reproduction in the Australian stingless bee Trigona carbonaria Smith (Hymenoptera, Apidae)

Abstract.Brood of the Australian stingless bee Trigona carbonaria was investigated using microsatellites to determine the origin of males. Genotypes of over 1800 males sampled from ten unrelated colonies were consistent with the hypothesis that the resident queen was the sole mother of the males and that workers, either natal or non-natal (parasitic), did not contribute to the production of males. Dissections of 300 workers from a total of four colonies showed that ovaries are present but not activated.

No evidence of queen thelytoky following interspecific crosses of the honey bees Apis cerana and Apis mellifera

The human-mediated dispersal of species over geographical boundaries can bring previously isolated sister taxa into contact. Interspecific mating between closely related species may then occur, with various outcomes ranging from hybridization to reproductive interference. In the case of the Eastern honey bee Apis cerana and the Western honey bee Apis mellifera, an additional possible reproductive outcome has been posited: interspecific sperm triggers queens to produce daughters from unfertilized eggs via thelytokous parthenogenesis. Such an outcome would go unnoticed in natural population mergers, as queens, which are polyandrous, are likely to mate with both conspecific and interspecific males. We performed reciprocal crosses between A. mellifera and A. cerana via artificial insemination, plus control inseminations of saline (five queens per species per treatment), and genetically assessed the sex and origin of any resulting offspring. Neither A. cerana nor A. mellifera queens produced viable female brood after receiving interspecific semen, indicating a high cost of interspecific mating for both species. In two A. cerana colonies headed by cross-inseminated queens, workers responded by activating ovaries and laying eggs that were mainly male but occasionally female (i.e. thelytokous, 2% of brood), despite the queen’s continued presence in the nest. We conclude that thelytoky is not a consistent response to interspecific mating by queens of A. mellifera or A. cerana. Rather, at least in A. cerana, when colonies are faced with “mis-mated” queens, it may be up to the workers to secure the reproductive future of the colony.