Fluid Nest Membership Drives Variable Relatedness in Groups of a Facultatively Social Bee

Kin selection theory has dominated our understanding of the evolution of group living. However, many animal groups form among non-relatives, which gain no indirect fitness benefits from cooperating with nestmates. In this study, we characterized the relatedness and inter-nest migration behavior of the facultatively social carpenter bee, Xylocopa sonorina. Nesting constraints due to costly nest construction in this species give rise to intense intraspecific competition over access to existing nests. We used mark-recapture techniques to characterize patterns of dispersal and nest relocation within a nesting aggregation of spatially clustered nests. Two-thirds of bees relocated at least once during the reproductive season, likely to seek reproductive opportunities in another nest. This fluid nest membership creates opportunities for association among non-relatives. To assess the effects of this dynamic nesting behavior on group relatedness, we used microsatellite analysis to estimate relative relatedness within and between nests in the aggregation. We found that relatedness was variable across sampling years, but that in many cases nestmates were no more related to one another than they were to non-nestmate bees in the population. Together, these results suggest that group composition in X. sonorina may result from strategies to maximize direct fitness. This study supports the hypothesis that factors beyond kinship, such as ecological constraints, are likely to drive group formation in this species.

Krüppel-homologue 1 Mediates Hormonally Regulated Dominance Rank in a Social Bee

Dominance hierarchies are ubiquitous in invertebrates and vertebrates, but little is known on how genes influence dominance rank. Our gaps in knowledge are specifically significant concerning female hierarchies, particularly in insects. To start filling these gaps, we studied the social bumble bee Bombus terrestris, in which social hierarchies among females are common and functionally significant. Dominance rank in this bee is influenced by multiple factors, including juvenile hormone (JH) that is a major gonadotropin in this species. We tested the hypothesis that the JH responsive transcription factor Krüppel homologue 1 (Kr-h1) mediates hormonal influences on dominance behavior. We first developed and validated a perfluorocarbon nanoparticles-based RNA interference protocol for knocking down Kr-h1 expression. We then used this procedure to show that Kr-h1 mediates the influence of JH, not only on oogenesis and wax production, but also on aggression and dominance rank. To the best of our knowledge, this is the first study causally linking a gene to dominance rank in social insects, and one of only a few such studies on insects or on female hierarchies. These findings are important for determining whether there are general molecular principles governing dominance rank across gender and taxa.

Social consequences of energetically costly nest construction in a facultatively social bee

Social groups form when the costs of breeding independently exceed fitness costs imposed by group living. The costs of independent breeding can often be energetic, especially for animals performing expensive behaviours, such as nest construction. To test the hypothesis that nesting costs can drive sociality by disincentivizing independent nest founding, we measured the energetics of nest construction and inheritance in a facultatively social carpenter bee ( Xylocopa sonorina Smith), which bores tunnel nests in wood. We measured metabolic rates of bees excavating wood and used computerized tomography images of nesting logs to measure excavation volumes. From these data, we demonstrate costly energetic investments in nest excavation of a minimum 4.3 kJ per offspring provisioned, an expense equivalent to nearly 7 h of flight. This high, potentially prohibitive cost of nest founding may explain why females compete for existing nests rather than constructing new ones, often leading to the formation of social groups. Further, we found that nest inheritors varied considerably in their investment in nest renovation, with costs ranging more than 12-fold (from 7.08 to 89.1 kJ energy), probably reflecting differences in inherited nest quality. On average, renovation costs were lower than estimated new nest construction costs, with some nests providing major savings. These results suggest that females may join social groups to avoid steep energetic costs, but that the benefits of this strategy are not experienced equally.

Bumblebees develop more efficient traplines than honey bees

Central place foraging pollinators, such as bees, tend to learn multi-destination routes (traplines) to efficiently visit known feeding locations and return to their nest. To what extent these routing behaviours are shared across species is unknown. Here we ran laboratory experiments to compare trapline formation and efficiency by foragers of two social bee species that differ in their collective foraging strategies: the solo foraging bumblebee Bombus terrestris and the mass foraging honey bee Apis mellifera. In a simple routing task with four artificial flowers, both bumblebees and honey bees developed a stable route, although honey bees were slower and less efficient to do so. In a more complex routing task with six flowers, only bumblebees developed a stable route. Honey bees took a longer time to discover all flowers and never integrated them in a single route. Simulations of a model of trapline formation show that these inter-specific differences can be replicated by adjusting the strength of a single learning parameter. Comparing bumblebees and honey bees in the same experimental conditions thus revealed key differences in their spatial foraging strategies, potentially driven by social constraints.

Neurotoxic alkaloid in pollen and nectar excludes generalist bees from foraging at death-camas, Toxicoscordion paniculatum (Melanthiaceae)

Many plants produce broadly active toxins to which specialist herbivores—typically insects—have evolved counter-adaptations, sometimes spawning co-evolutionary arms races. Many non-social bee species are likewise taxonomic host specialists, but the specialists’ pollen hosts frequently attract diverse floral generalists as well, even to flowers of plants that are otherwise chemically defended. In this study of foothills death-camas, Toxicoscordion paniculatum (Nutt.) Rydberg (formerly Zigadenus), we show that its pollen and nectar both contain zygacine, the steroidal alkaloid responsible for this plant’s notorious mammalian toxicity. Hungry naïve adults of a generalist solitary bee, Osmia lignaria Say (Megachilidae), would briefly drink death-camas nectar or biologically relevant doses of zygacine in syrup, followed by prolonged bouts of irritable tongue grooming; many became paralyzed and some even died. Larvae fed dosed provision masses likewise often ceased feeding and sometimes died. Prolonged irritation and subsequent deterrence of foraging O. lignaria likely illustrates why it and 50+ other vernal bee species were absent from death-camas flowers in a five-state survey. The sole visiting bee, Andrena astragali, foraged exclusively at death-camas flowers for pollen and nectar. Thus, a toxic alkaloid found in death-camas pollen and nectar deters generalist bees from flowers of this pollinator-dependent monocot, restricting visitation to a single specialist bee that tolerates death-camas toxins and is its likely pollinator.

Reduced Water Negatively Impacts Social Bee Survival and Productivity Via Shifts in Floral Nutrition

Pollinators provide a key ecosystem service vital for the survival and stability of the biosphere. Identifying factors influencing the plant–pollinator mutualism and pollinator management is necessary for maintaining a healthy ecosystem. Since healthy beehives require substantial amounts of carbohydrates (nectar) and protein (pollen) from forage plants such as clover, we must assess how resources offered by plants change under limited water conditions in order to fully understand how drought modifies the pollination mutualism. Here we document how reduced water availability leads to decreased nectar quality and quantity and decreased protein quality of pollen. Furthermore, we provide conclusive evidence that these lower quality resources lead to decreased survival and productivity in both developing honey bees (Hymenoptera: Apidae) and bumble bees (Hymenoptera: Apidae). The results emphasize the importance of the nutritional effects of reduced water on bees when predicting shifts of pollination mutualisms under climate change.

Unrelated males in colonies of facultatively social bee

Colonies of social Hymenoptera are usually groups of closely related females, in which the dominant female(s) is specialized for reproduction and subordinate females care for immature offspring. Kin selection is thought to be the main factor that supports social cohesion. We have discovered a simple society of the bee Ceratina chalybea with an average of 4.68 colony members that cannot be maintained by kin selection alone. These colonies consisted of old reproductive female, young adults and provisioned brood cells. About half of young adults are unrelated to the old female, and almost all of the young adults are male. The old female provisions new brood cells, while continuing to feed young adult offspring. As young adults do not perform demanding or risky activities, they incur little or no cost, but they do benefit from the food they obtain from the old female.