Controlling the Uptake and Release of Semiochemicals in Channel-Type Metal-Organic Frameworks Through Pore Expansion.

Semiochemicals can be used to manipulate insect behaviour for sustainable pest management strategies, but their high volatility is a major issue for their practical implementation. Inclusion of these molecules within porous materials is a potential solution to this issue, as it can allow for a slower and more controlled release. In this work, we demonstrate that a series of Zr(IV) and Al(III) metal-organic frameworks (MOFs) with channel-type pores enable controlled release of three semiochemicals over 100 days by pore size design, with the uptake and rate of release highly dependent on the pore size. Insight from grand canonical Monte Carlo simulations indicates that this is due to weaker MOF-guest interactions per guest molecule as the pore size increases. These MOFs are all stable post-release and can be reloaded to show near-identical re-release profiles. These results provide valuable insight on the diffusion behaviour of volatile guests in MOFs, and for the further development of porous materials for sustainable agriculture applications.

Limited introgression from non-native commercial strains and signatures of adaptation in the key pollinator Bombus terrestris.

Insect pollination is fundamental for natural ecosystems and agricultural crops. The bumblebee species Bombus terrestris has become a popular choice for commercial crop pollination worldwide due to its effectiveness and ease of mass rearing. Bumblebee colonies are mass produced for the pollination of more than 20 crops and imported into over 50 countries including countries outside their native ranges, and the risk of invasion by commercial non-native bumblebees is considered an emerging issue for global conservation and biological diversity. Here, we use genome-wide data from seven wild populations close to and far from farms using commercial colonies, as well as commercial populations, to investigate the implications of utilizing commercial bumblebee subspecies in the UK. We find evidence for generally low levels of introgression between commercial and wild bees, with higher admixture proportions in the bees occurring close to farms. We identify genomic regions putatively involved in local and global adaptation, and genes in locally adaptive regions were found to be enriched for functions related to taste receptor activity, oxidoreductase activity, fatty acid and lipid biosynthetic processes. Despite more than 30 years of bumblebee colony importation into the UK, we observe low impact on the genetic integrity of local B. terrestris populations, but we highlight that even limited introgression might negatively affect locally adapted populations.

Multiple stressors interact to impair the performance of bumblebee Bombus terrestris colonies.

Bumblebees are constantly exposed to a wide range of biotic and abiotic stresses which they must defend themselves against to survive. Pathogens and pesticides represent important stressors that influence bumblebee health, both when acting alone or in combination. To better understand bumblebee health, we need to investigate how these factors interact, yet experimental studies to date generally focus on only one or two stressors. The aim of this study is to evaluate how combined effects of four important stressors (the gut parasite Nosema ceranae, the neonicotinoid insecticide thiamethoxam, the pyrethroid insecticide cypermethrin and the EBI fungicide tebuconazole) interact to affect bumblebees at the individual and colony levels. We established seven treatment groups of colonies that we pulse exposed to different combinations of these stressors for 2 weeks under laboratory conditions. Colonies were subsequently placed in the field for 7 weeks to evaluate the effect of treatments on the prevalence of N. ceranae in inoculated bumblebees, expression levels of immunity and detoxification-related genes, food collection, weight gain, worker and male numbers, and production of worker brood and reproductives. Exposure to pesticide mixtures reduced food collection by bumblebees. All immunity-related genes were upregulated in the bumblebees inoculated with N. ceranae when they had not been exposed to pesticide mixtures, and bumblebees exposed to the fungicide and the pyrethroid were less likely to have N. ceranae. Combined exposure to the three-pesticide mixture and N. ceranae reduced bumblebee colony growth, and all treatments had detrimental effects on brood production. The groups exposed to the neonicotinoid insecticide produced 40%-76% fewer queens than control colonies. Our findings show that exposure to combinations of stressors that bumblebees frequently come into contact with have detrimental effects on colony health and performance and could therefore have an impact at the population level. These results also have significant implications for current practices and policies for pesticide risk assessment and use as the combinations tested here are frequently applied simultaneously in the field. Understanding the interactions between different stressors will be crucial for improving our ability to manage bee populations and for ensuring pollination services into the future.

Los abejorros están constantemente expuestos a una amplia gama de agentes estresantes bióticos y abióticos de los que deben defenderse para sobrevivir. Los patógenos y los pesticidas son importantes factores estresantes que influyen en la salud de los abejorros, tanto cuando actúan solos como en combinación. Para tener un mejor conocimiento sobre la salud de los abejorros, debemos investigar cómo interactúan estos factores estresantes, pero los estudios experimentales hasta la fecha generalmente se centran en estudiar solo uno o dos factores. El objetivo de nuestro estudio es evaluar cómo los efectos combinados de cuatro importantes factores estresantes (el parásito intestinal Nosema ceranae, el insecticida neonicotinoide tiametoxam, el insecticida piretroide cipermetrina y el fungicida EBI tebuconazol) interactúan para afectar a los abejorros a nivel individual y de colonia. Establecimos siete grupos de tratamiento de colonias de abejorros que expusimos a diferentes combinaciones de estos factores estresantes durante dos semanas en condiciones de laboratorio, y posteriormente se colocaron en el campo durante siete semanas, para evaluar el efecto de los tratamientos sobre la prevalencia de N. ceranae en abejorros inoculados, los niveles de expresión de genes relacionados con la inmunidad y la desintoxicación, la recolección de alimentos, el aumento de peso, el número de obreras y machos, y la producción de cría de obreras, machos y reinas. La exposición a mezclas de pesticidas redujo la recolección de alimentos por parte de los abejorros. Todos los genes relacionados con la inmunidad se sobre-expresaron en los abejorros inoculados con N. ceranae cuando no habían estado expuestos a mezclas de pesticidas, y los abejorros expuestos al fungicida y al piretroide presentaron menos probabilidades de tener N. ceranae. La exposición combinada a la mezcla de tres pesticidas y N. ceranae redujo el crecimiento de la colonia de abejorros y todos los tratamientos tuvieron efectos perjudiciales en la producción de crías. Los grupos expuestos al insecticida neonicotinoide produjeron entre un 40 y un 76% menos de reinas que las colonias control. Nuestros hallazgos muestran que la exposición a combinaciones de factores estresantes con los que los abejorros entran frecuentemente en contacto tiene efectos perjudiciales sobre la salud y el rendimiento de la colonia y, por lo tanto, podría tener un impacto a nivel poblacional. Estos resultados también tienen importantes implicaciones para las prácticas y políticas actuales de evaluación de riesgos y uso de plaguicidas, ya que las combinaciones probadas aquí se aplican con frecuencia simultáneamente en el campo. Comprender las interacciones entre los diferentes factores de estrés es fundamental para mejorar nuestra capacidad de gestión de las poblaciones de abejas y así garantizar los servicios de polinización en el futuro.

Limited female dispersal predicts the incidence of Wolbachia across ants (Hymenoptera: Formicidae).

The endosymbiotic bacterium Wolbachia is perhaps the greatest panzootic in the history of life on Earth, yet remarkably little is known regarding the factors that determine its incidence across species. One possibility is that Wolbachia more easily invades species with structured populations, due to the increased strength of genetic drift and higher initial frequency of infection. This should enable strains that induce mating incompatibilities to more easily cross the threshold prevalence above which they spread to either fixation or a stable equilibrium infection prevalence. Here, we provide empirical support for this hypothesis by analysing the relationship between female dispersal (as a proxy for population structure) and the incidence of Wolbachia across 250 species of ants. We show that species in which the dispersal of reproductive females is limited are significantly more likely to be infected with Wolbachia than species whose reproductive ecology is consistent with significant dispersal of females, and that this relationship remains after controlling for host phylogeny. We suggest that structured host populations, in this case resulting from limited female dispersal, may be an important feature determining how easily Wolbachia becomes successfully established in a novel host, and thus its occurrence across a wide diversity of invertebrate hosts.

Molecular signatures of plastic phenotypes in two eusocial insect species with simple societies.

Phenotypic plasticity is important in adaptation and shapes the evolution of organisms. However, we understand little about what aspects of the genome are important in facilitating plasticity. Eusocial insect societies produce plastic phenotypes from the same genome, as reproductives (queens) and nonreproductives (workers). The greatest plasticity is found in the simple eusocial insect societies in which individuals retain the ability to switch between reproductive and nonreproductive phenotypes as adults. We lack comprehensive data on the molecular basis of plastic phenotypes. Here, we sequenced genomes, microRNAs (miRNAs), and multiple transcriptomes and methylomes from individual brains in a wasp (Polistes canadensis) and an ant (Dinoponera quadriceps) that live in simple eusocial societies. In both species, we found few differences between phenotypes at the transcriptional level, with little functional specialization, and no evidence that phenotype-specific gene expression is driven by DNA methylation or miRNAs. Instead, phenotypic differentiation was defined more subtly by nonrandom transcriptional network organization, with roles in these networks for both conserved and taxon-restricted genes. The general lack of highly methylated regions or methylome patterning in both species may be an important mechanism for achieving plasticity among phenotypes during adulthood. These findings define previously unidentified hypotheses on the genomic processes that facilitate plasticity and suggest that the molecular hallmarks of social behavior are likely to differ with the level of social complexity.

Alarm Pheromone Composition and Behavioral Activity in Fungus-Growing Ants.

Chemical communication is a dominant method of communication throughout the animal kingdom and can be especially important in group-living animals in which communicating threats, either from predation or other dangers, can have large impacts on group survival. Social insects, in particular, have evolved a number of pheromonal compounds specifically to signal alarm. There is predicted to be little selection for interspecific variation in alarm cues because individuals may benefit from recognizing interspecific as well as conspecific cues and, consequently, alarm cues are not normally thought to be used for species or nestmate recognition. Here, we examine the composition of the alarm pheromones of seven species of fungus-growing ants (Attini), including both basal and derived species and examine the behavioral responses to alarm pheromone of Acromyrmex leaf-cutting ants, the sister genus to the highly studied Atta leaf-cutting ants. We find surprisingly high interspecific variation in alarm pheromone composition across the attine phylogeny. Interestingly, the active component of the alarm pheromone was different between the two leaf-cutting ant genera. Furthermore, in contrast to previous studies on Atta, we found no differences among morphological castes in their responses to alarm pheromone in Acromyrmex but we did find differences in responses among putative age classes. The results suggest that the evolution of alarm communication and signaling within social insect clades can be unexpectedly complex and that further work is warranted to understand whether the evolution of different alarm pheromone compounds is adaptive.

Only full-sibling families evolved eusociality.

Arising from M. A. Nowak, C. E. Tarnita & E. O. Wilson 466, 1057-1062 (2010); Nowak et al. reply. The paper by Nowak et al. has the evolution of eusociality as its title, but it is mostly about something else. It argues against inclusive fitness theory and offers an alternative modelling approach that is claimed to be more fundamental and general, but which, we believe, has no practical biological meaning for the evolution of eusociality. Nowak et al. overlook the robust empirical observation that eusociality has only arisen in clades where mothers are associated with their full-sibling offspring; that is, in families where the average relatedness of offspring to siblings is as high as to their own offspring, independent of population structure or ploidy. We believe that this omission makes the paper largely irrelevant for understanding the evolution of eusociality.

Pleiotropic effects of juvenile hormone in ant queens and the escape from the reproduction-immunocompetence trade-off.

The ubiquitous trade-off between survival and costly reproduction is one of the most fundamental constraints governing life-history evolution. In numerous animals, gonadotropic hormones antagonistically suppressing immunocompetence cause this trade-off. The queens of many social insects defy the reproduction-survival trade-off, achieving both an extraordinarily long life and high reproductive output, but how they achieve this is unknown. Here we show experimentally, by integrating quantification of gene expression, physiology and behaviour, that the long-lived queens of the ant Lasius niger have escaped the reproduction-immunocompetence trade-off by decoupling the effects of a key endocrine regulator of fertility and immunocompetence in solitary insects, juvenile hormone (JH). This modification of the regulatory architecture enables queens to sustain a high reproductive output without elevated JH titres and suppressed immunocompetence, providing an escape from the reproduction-immunocompetence trade-off that may contribute to the extraordinary lifespan of many social insect queens.

Behavioural effects of juvenile hormone and their influence on division of labour in leaf-cutting ant societies.

Division of labour in social insects represents a major evolutionary transition, but the physiological mechanisms that regulate this are still little understood. Experimental work with honey bees, and correlational analyses in other social insects, have implicated juvenile hormone (JH) as a regulatory factor, but direct experimental evidence of behavioural effects of JH in social insects is generally lacking. Here, we used experimental manipulation of JH to show that raised JH levels in leaf-cutting ants results in workers becoming more active, phototactic and threat responsive, and engaging in more extranidal activity - behavioural changes that we show are all characteristic of the transition from intranidal work to foraging. These behavioural effects on division of labour suggest that the JH mediation of behaviour occurs across multiple independent evolutions of eusociality, and may be a key endocrine regulator of the division of labour which has produced the remarkable ecological and evolutionary success of social insects.

Cryptic lineages hybridize for worker production in the harvester ant Messor barbarus.

The reproductive division of labour between queen and worker castes in social insects is a defining characteristic of eusociality and a classic example of phenotypic plasticity. Whether social insect larvae develop into queens or workers has long been thought to be determined by environmental cues, i.e. larvae are developmentally totipotent. Contrary to this paradigm, several recent studies have revealed that caste is determined by genotype in some ant species, but whether this is restricted to just a few exceptional species is still unclear. Here, we show that the Mediterranean harvester ant Messor barbarus possesses an unusual reproductive system, in which the female castes are genetically determined. Using both nuclear and mitochondrial data, we show that Iberian populations have two distinct, cryptic lineages. Workers are always inter-lineage hybrids whereas queens are always produced from pure-lineage matings. The results suggest that genetic caste determination may be more widespread in ants than previously thought, and that further investigation in other species is needed to understand the frequency and evolution of this remarkable reproductive system.

The effects of single and mixed infections of Apicystis bombi and deformed wing virus in Bombus terrestris.

Many pollinators are currently suffering from declines, diminishing their gene pool and increasing their vulnerability to parasites. Recently, an increasing diversity of parasites has been recorded in bumblebees, yet for many, knowledge of their virulence and hence the risk their presence poses, is lacking. The deformed wing virus (DWV), known to be ubiquitous in honey bees, has now been detected in bumblebees. In addition, the neogregarine Apicystis bombi has been discovered to be more prevalent than previously thought. Here, we assess for the first time the lethal and sublethal effects of these parasites during single and mixed infections of worker bumblebees (Bombus terrestris). Fifteen days after experimental exposure, 22% of bees exposed to A. bombi, 50% of bees exposed to DWV and 86% of bees exposed to both parasites had died. Bumblebees that had ingested A. bombi had increased sucrose sensitivity (SS) and a lower lipid:body size ratio than control bees. While dual infected bumblebees showed no increase in SS. Overall, we find that A. bombi exhibits both lethal and sublethal effects. DWV causes lethal effect and may reduce the sub lethal effects imposed by A. bombi. The results show that both parasites have significant, negative effects on bumblebee health, making them potentially of conservation concern.

Parasites in bloom: flowers aid dispersal and transmission of pollinator parasites within and between bee species.

The dispersal of parasites is critical for epidemiology, and the interspecific vectoring of parasites when species share resources may play an underappreciated role in parasite dispersal. One of the best examples of such a situation is the shared use of flowers by pollinators, but the importance of flowers and interspecific vectoring in the dispersal of pollinator parasites is poorly understood and frequently overlooked. Here, we use an experimental approach to show that during even short foraging periods of 3 h, three bumblebee parasites and two honeybee parasites were dispersed effectively onto flowers by their hosts, and then vectored readily between flowers by non-host pollinator species. The results suggest that flowers are likely to be hotspots for the transmission of pollinator parasites and that considering potential vector, as well as host, species will be of general importance for understanding the distribution and transmission of parasites in the environment and between pollinators.

Targeted agri-environment schemes significantly improve the population size of common farmland bumblebee species.

Changes in agricultural practice across Europe and North America have been associated with range contractions and local extinction of bumblebees (Bombus spp.). A number of agri-environment schemes have been implemented to halt and reverse these declines, predominantly revolving around the provision of additional forage plants. Although it has been demonstrated that these schemes can attract substantial numbers of foraging bumblebees, it remains unclear to what extent they actually increase bumblebee populations. We used standardized transect walks and molecular techniques to compare the size of bumblebee populations between Higher Level Stewardship (HLS) farms implementing pollinator-friendly schemes and Entry Level Stewardship (ELS) control farms. Bumblebee abundance on the transect walks was significantly higher on HLS farms than ELS farms. Molecular analysis suggested maximum foraging ranges of 566 m for Bombus hortorum, 714 m for B. lapidarius, 363 m for B. pascuorum and 799 m for B. terrestris. Substantial differences in maximum foraging range were found within bumblebee species between farm types. Accounting for foraging range differences, B. hortorum (47 vs 13 nests/km(2) ) and B. lapidarius (45 vs 22 nests/km(2) ) were found to nest at significantly greater densities on HLS farms than ELS farms. There were no significant differences between farm type for B. terrestris (88 vs 38 nests/km(2) ) and B. pascuorum (32 vs 39 nests/km(2) ). Across all bumblebee species, HLS management had a significantly positive effect on bumblebee nest density. These results show that targeted agri-environment schemes that increase the availability of suitable forage can significantly increase the size of wild bumblebee populations.

Immunosenescence and resistance to parasite infection in the honey bee, Apis mellifera.

Immunosenescence, the systemic reduction of immune efficiency with age, is increasingly recognised as having important implications for host-parasite dynamics. Changes in the immune response can impact on the ability of an individual to resist or moderate parasite infection, depending on how and when it encounters a parasite challenge. Using the European honey bee Apis mellifera and its microsporidian parasite Nosema ceranae, we investigated the effects of host age on the ability to resist parasite infection and on baseline immunocompetence, assessed by quantifying constitutive (PO) and potential levels (PPO) of the phenoloxidase immune enzyme as general measures of immune function. There was a significant correlation between the level of general immune function and infection intensity, but not with survival, and changes in immune function with age correlated with the ability of individuals to resist parasite infection. Older individuals had better survival when challenged with a parasite than younger individuals, however they also had more intense infections and lower baseline immunocomptence. The ability of older individuals to have high infection intensities yet live longer, has potential consequences for parasite transmission. The results highlight the need to consider age in host-parasite studies and show the importance of choosing the correct measure when assaying invertebrate immunity.

Identification of fungi isolated from commercial bumblebee colonies.

Fungi can have important beneficial and detrimental effects on animals, yet our understanding of the diversity and function of most bee-associated fungi is poor. Over 2 million bumblebee colonies are traded globally every year, but the presence and transport of viable fungi within them is unknown. Here, we explored whether any culturable fungi could be isolated from commercial bumblebee nests. We collected samples of various substrates from within 14 bumblebee colonies, including the honey, honey cup wall, egg cup wall, and frass then placed them on agar and recorded any growth. Fungal morphotypes were then subcultured and their ITS region sequenced for identification. Overall, we cultured 11 fungal species from the various nest substrates. These included both pathogenic and non-pathogenic fungi, such as Aspergillus sp., Penicillium sp., and Candida sp. Our results provide the first insights into the diversity of viable fungal communities in commercial bumblebee nests. Further research is needed to determine if these fungi are unique to commercial colonies or prevalent in wild bumblebee nests, and crucially to determine the ecological and evolutionary implications of these fungi in host colonies.

Biogenic amines are associated with worker task but not patriline in the leaf-cutting ant Acromyrmex echinatior.

Division of labor among eusocial insect workers is a hallmark of advanced social organization, but its underlying neural mechanisms are not well understood. We investigated whether differences in whole-brain levels of the biogenic amines dopamine (DA), serotonin (5HT), and octopamine (OA) are associated with task specialization and genotype in similarly sized and aged workers of the leaf-cutting ant Acromyrmex echinatior, a polyandrous species in which genotype correlates with worker task specialization. We compared amine levels of foragers and waste management workers to test for an association with worker task, and young in-nest workers across patrilines to test for a genetic influence on brain amine levels. Foragers had higher levels of DA and OA and a higher OA:5HT ratio than waste management workers. Patrilines did not significantly differ in amine levels or their ratios, although patriline affected worker body size, which correlated with amine levels despite the small size range sampled. Levels of all three amines were correlated within individuals in both studies. Among patrilines, mean levels of DA and OA, and OA and 5HT were also correlated. Our results suggest that differences in biogenic amines could regulate worker task specialization, but may be not be significantly affected by genotype.

Tolerating an infection: an indirect benefit of co-founding queen associations in the ant Lasius niger.

Pathogens exert a strong selection pressure on organisms to evolve effective immune defences. In addition to individual immunity, social organisms can act cooperatively to produce collective defences. In many ant species, queens have the option to found a colony alone or in groups with other, often unrelated, conspecifics. These associations are transient, usually lasting only as long as each queen benefits from the presence of others. In fact, once the first workers emerge, queens fight to the death for dominance. One potential advantage of co-founding may be that queens benefit from collective disease defences, such as mutual grooming, that act against common soil pathogens. We test this hypothesis by exposing single and co-founding queens to a fungal parasite, in order to assess whether queens in co-founding associations have improved survival. Surprisingly, co-foundresses exposed to the entomopathogenic fungus Metarhizium did not engage in cooperative disease defences, and consequently, we find no direct benefit of multiple queens on survival. However, an indirect benefit was observed, with parasite-exposed queens producing more brood when they co-founded, than when they were alone. We suggest this is due to a trade-off between reproduction and immunity. Additionally, we report an extraordinary ability of the queens to tolerate an infection for long periods after parasite exposure. Our study suggests that there are no social immunity benefits for co-founding ant queens, but that in parasite-rich environments, the presence of additional queens may nevertheless improve the chances of colony founding success.

Emerging dangers: deadly effects of an emergent parasite in a new pollinator host.

There is growing concern about the threats facing many pollinator populations. Emergent diseases are one of the major threats to biodiversity and a microsporidian parasite, Nosema ceranae, has recently jumped host from the Asian to the Western honeybee, spreading rapidly worldwide, and contributing to dramatic colony losses. Bumblebees are ecologically and economically important pollinators of conservation concern, which are likely exposed to N. ceranae by sharing flowers with honeybees. Whilst a further intergeneric jump by N. ceranae to infect bumblebees would be potentially serious, its capacity to do this is unknown. Here we investigate the prevalence of N. ceranae in wild bumblebees in the UK and determine the infectivity of the parasite under controlled conditions. We found N. ceranae in all seven wild bumblebee species sampled, and at multiple sites, with many of the bees having spores from this parasite in their guts. When we fed N. ceranae spores to bumblebees under controlled conditions, we confirmed that the parasite can infect bumblebees. Infections spread from the midgut to other tissues, reduced bumblebee survival by 48% and had sub-lethal effects on behaviour. Although spore production appeared lower in bumblebees than in honeybees, virulence was greater. The parasite N. ceranae therefore represents a real and emerging threat to bumblebees, with the potential to have devastating consequences for their already vulnerable populations.

Nutritional limitation and resistance to opportunistic Aspergillus parasites in honey bee larvae.

Honey bees are threatened by land use changes which reduce the availability and diversity of pollen and nectar resources. There is concern that poor nutrition may be involved in recent population declines, either directly or due to indirect effects on immunocompetence. The larval stage is likely to be the most vulnerable to a poor diet, but the effects of larval nutrition on the disease susceptibility of bees are not well known. In this study we used laboratory-reared honey bee larvae to investigate the effects of diet quality on disease susceptibility to the opportunistic fungal parasites Aspergillus flavus, Aspergillus phoenicis and A. fumigatus. Larvae fed on a nutritionally poor diet were found to be significantly more susceptible to A. fumigatus. Larval resistance to A. fumigatus was enhanced by feeding with a diet supplemented with either dandelion or polyfloral pollens. This indicates that dandelion and polyfloral pollens contain elements that enhance resistance to this fungal disease, illustrating an interaction between nutrition and parasitism and emphasising the benefit of diverse floral resources in the environment to maintain honey bee health.

Kin-informative recognition cues in ants.

Although social groups are characterized by cooperation, they are also often the scene of conflict. In non-clonal systems, the reproductive interests of group members will differ and individuals may benefit by exploiting the cooperative efforts of other group members. However, such selfish behaviour is thought to be rare in one of the classic examples of cooperation--social insect colonies--because the colony-level costs of individual selfishness select against cues that would allow workers to recognize their closest relatives. In accord with this, previous studies of wasps and ants have found little or no kin information in recognition cues. Here, we test the hypothesis that social insects do not have kin-informative recognition cues by investigating the recognition cues and relatedness of workers from four colonies of the ant Acromyrmex octospinosus. Contrary to the theoretical prediction, we show that the cuticular hydrocarbons of ant workers in all four colonies are informative enough to allow full-sisters to be distinguished from half-sisters with a high accuracy. These results contradict the hypothesis of non-heritable recognition cues and suggest that there is more potential for within-colony conflicts in genetically diverse societies than previously thought.