Social insect colonies exhibit a variety of life history strategies, from the annual, semelparous colonies of temperate bees and wasps to the long-lived colonies of many ants and honeybees. Species introduced to novel habitats may exhibit plasticity in life history strategies as a result of the introduction, but the factors governing these changes often remain obscure. Vespula pensylvanica, a yellowjacket wasp, exhibits such plasticity in colony longevity. Multi-year (perennial) colonies are relatively common in introduced populations in Hawaii, while source populations in the western United States are typically on an annual cycle. Here, we use experiments and observational data to examine how diet, disease, nest thermal environment, and nest location influence colony longevity in a population with both annual and perennial colonies. Counter to our predictions, experimental feeding and warming did not increase colony survival in the winter in the introduced range. However, Moku Virus load and wasp colony density predicted colony survival in one year, suggesting a potential role for disease in modulating colony phenology. We also found that local V. pensylvanica colony density was positively correlated with Moku Virus loads, and that Arsenophonus sp. bacterial loads in V. pensylvanica colonies were positively associated with proximity to feral honeybee (Apis mellifera) hives, suggesting potential transmission routes for these poorly understood symbionts. The factors influencing colony longevity in this population are likely multiple and interactive. More important than food availability, we propose winter precipitation as a critical factor that may explain temporal and spatial variation in colony longevity in these invasive wasps.

Invasive species present a worldwide concern as competition and pathogen reservoirs for native species. Specifically, the invasive social wasp, Vespula pensylvanica, is native to western North America and has become naturalized in Hawaii, where it exerts pressures on native arthropod communities as a competitor and predator. As invasive species may alter the microbial and disease ecology of their introduced ranges, there is a need to understand the microbiomes and virology of social wasps. We used 16S rRNA gene sequencing to characterize the microbiome of V. pensylvanica samples pooled by colony across two geographically distinct ranges and found that wasps generally associate with taxa within the bacterial genera Fructobacillus, Fructilactobacillus, Lactococcus, Leuconostoc, and Zymobacter, and likely associate with environmentally-acquired bacteria. Furthermore, V. pensylvanica harbors-and in some cases were dominated by-many endosymbionts including Wolbachia, Sodalis, Arsenophonus, and Rickettsia, and were found to contain bee-associated taxa, likely due to scavenging on or predation upon honey bees. Next, we used reverse-transcriptase quantitative PCR to assay colony-level infection intensity for Moku virus (family: Iflaviridae), a recently-described disease that is known to infect multiple Hymenopteran species. While Moku virus was prevalent and in high titer, it did not associate with microbial diversity, indicating that the microbiome may not directly interact with Moku virus in V. pensylvanica in meaningful ways. Collectively, our results suggest that the invasive social wasp V. pensylvanica associates with a simple microbiome, may be infected with putative endosymbionts, likely acquires bacterial taxa from the environment and diet, and is often infected with Moku virus. Our results suggest that V. pensylvanica, like other invasive social insects, has the potential to act as a reservoir for bacteria pathogenic to other pollinators, though this requires experimental demonstration.

Emerging infectious diseases (EIDs) are a global threat to honeybees, and spillover from managed bees threaten wider insect populations. Deformed wing virus (DWV), a widespread virus that has become emergent in conjunction with the spread of the mite Varroa destructor, is thought to be partly responsible for global colony losses. The arrival of Varroa in honeybee populations causes a dramatic loss of viral genotypic diversity, favouring a few virulent strains. Here, we investigate DWV spillover in an invasive Hawaiian population of the wasp, Vespula pensylvanica, a honeybee predator and honey-raider. We show that Vespula underwent a parallel loss in DWV variant diversity upon the arrival of Varroa, despite the mite being a honeybee specialist. The observed shift in Vespula DWV and the variant-sharing between Vespula and Apis suggest that these wasps can acquire DWV directly or indirectly from honeybees. Apis prey items collected from Vespula foragers were positive for DWV, indicating predation is a possible route of transmission. We also sought cascading effects of DWV shifts in a broader Vespula pathogen community. We identified concurrent changes in a suite of additional pathogens, as well as shifts in the associations between these pathogens in Vespula. These findings reveal how hidden effects of the Varroa mite can, via spillover, transform the composition of pathogens in interacting species, with potential knock-on effects for entire pathogen communities.

Invasive social insects rank among the most damaging of terrestrial species. They are responsible for extensive damage and severely threaten the biodiversity of environments where they are introduced. Variation in colony social form commonly occurs in introduced populations of yellowjacket wasps (genus Vespula). In particular, invasive colonies may contain multiple queens (i.e., polygyne) and persist several years, while in the native range, the colonies are usually annual and harbor a single queen (i.e., monogyne). In this study, we used genome-wide loci obtained by double digest restriction site-associated DNA sequencing (RADseq) to investigate the genetic structure and queen turnover in colonies of the western yellowjacket, Vespula pensylvanica, in their introduced range in Hawaii. Of the 27 colonies monitored over four months (October-January), 19 were polygyne and already contained multiple queens on the first day of sampling. Contrary to previous speculation, this finding suggests that polygyny often arises early in the annual colony cycle, before the production of new queens in the fall. Furthermore, polygyne colonies exhibited a prolonged average lifespan relative to those headed by a single queen. As a result, there is no clear window during which colony eradication efforts would be more effective than upon first discovery. The relatedness among nestmate queens was slightly above zero, indicating that these colonies are generally composed of nonrelatives. The queen turnover within each colony was low, and we detected some full-sibling workers sampled up to four months apart. Finally, we did not detect any population structure among colonies, suggesting that queens disperse up to several kilometers. Taken together, our results provide the first insights into the requeening dynamics in this invasive and incipiently polygyne population and illuminate the early establishment of multiple long-lasting queens in these damaging colonies.

Cooperative breeding decreases the direct reproductive output of subordinate individuals, but cooperation can be evolutionarily favored when there are challenges or constraints to breeding independently. Environmental factors, including temperature, precipitation, latitude, high seasonality, and environmental harshness have been hypothesized to correlate with the presence of cooperative breeding. However, to test the relationship between cooperation and ecological constraints requires comparative data on the frequency and variation of cooperative breeding across differing environments, ideally replicated across multiple species. Paper wasps are primitively social species, forming colonies composed of reproductively active dominants and foraging subordinates. Adult female wasps, referred to as foundresses, initiate new colonies. Nests can be formed by a single solitary foundress (noncooperative) or by multiple foundress associations (cooperative). Cooperative behavior varies within and among species, making paper wasps species well suited to disentangling ecological correlates of variation in cooperative behavior. This data set reports the frequency and extent of cooperative nest founding for 87 paper wasp species. Data were assembled from more than 170 published sources, previously unpublished field observations, and photographs contributed by citizen scientists to online natural history repositories. The data set includes 25,872 nest observations and reports the cooperative behavioral decisions for 45,297 foundresses. Species names were updated to reflect modern taxonomic revisions. The type of substrate on which the nest was built is also included, when available. A smaller population-level version of this data set found that the presence or absence of cooperative nesting in paper wasps was correlated with temperature stability and environmental harshness, but these variables did not predict the extent of cooperation within species. This expanded data set contains details about individual nests and further increases the power to address the relationship between the environment and the presence and extent of cooperative breeding. Beyond the ecological drivers of cooperation, these high-resolution data will be useful for future studies examining the evolutionary consequences of variation in social behavior. This data set may be used for research or educational purposes provided that this data paper is cited.