Ant Formica Fusca Research Articles

Queen fecundity, worker entourage and cuticular chemistry in the ant Formica fusca.

Cooperative breeding entails conflicts over reproductive shares that may be settled in different ways. In ants, where several queens simultaneously reproduce in a colony, both queens and workers may influence the reproductive apportionment and offspring quality. Queens may vary in their intrinsic fecundity, which may influence the size of the worker entourage attending individual queens, and this may eventually dictate the reproductive output of a queen. We tested whether the reproductive success of queens is affected by the size of their worker entourage, their fecundity at the onset of the reproductive season, and whether the queen cuticular hydrocarbon profile carries information on fecundity. We show that in the ant Formica fusca both queen fecundity and egg hatching success increase with the size of their entourage, and that newly hatched larvae produced by initially highly fecund queens are smaller. Furthermore, higher relatedness among workers increased queen fecundity. Finally, the queens that received a large worker entourage differed in the cuticular chemistry from those that received a small worker entourage. Our results thus show that workers play a pivotal role in determining queen fitness, that high intracolony relatedness among workers enhances the overall reproductive output in the colony, and that queen fecundity is reflected in their cuticular hydrocarbon profile.

Relatedness modulates reproductive competition among queens in ant societies with multiple queens.

Reproductive sharing in animal groups with multiple breeders, insects and vertebrates alike, contains elements of both conflict and cooperation, and depends on both relatedness between co-breeders, as well as their internal and external conditions. We studied how queens of the ant Formica fusca adjust their reproductive efforts in response to experimental manipulations of the kin competition regime in their nest. Queens respond to the presence of competitors by increasing their egg laying efforts, but only if the competitors are highly fecund and distantly related. Such a mechanism is likely to decrease harmful competition among close relatives. We demonstrate that queens of Formica fusca fine-tune their cooperative breeding behaviors in response to kinship and fecundity of others in a remarkably precise and flexible manner.

Similarities in Recognition Cues Lead to the Infiltration of Non-Nestmates in an Ant Species.

Chemical cues are among the most important information-sharing mechanisms in insect societies, in which cuticular hydrocarbons play a central role, e.g., from nestmate recognition to queen signaling. The nestmate recognition mechanism usually prevents intruders from taking advantage of the resources stored in the nest. However, nestmate recognition is not unconditionally effective, and foreign individuals can sometimes infiltrate unrelated nests and take advantage of the colony resources. In this study, we investigated the role of overall colony odor profiles on the ability of conspecific workers to drift into unrelated colonies. We hypothesized that drifters would have higher chances of success by infiltrating colonies with the odor profiles most similar to their own nest, avoiding being detected as non-nestmates. By performing a drifting bioassay, we found that workers of the ant Formica fusca infiltrated unrelated conspecific colonies at a rate of 2.4%, significantly infiltrating colonies displaying CHC profiles most similar to their natal nests. Notably, methyl branched hydrocarbons seem to play a role as recognition cues in this species. In addition, we show that environmental rather than genetic factors are responsible for most contributions on the CHC phenotype, presenting ca. of 50% and 27.5% of explained variation respectively, and playing a major role in how worker ants detect and prevent the infiltration of non-nestmates in the colony. Hence, relying on cuticular hydrocarbons similarities could be a profitably evolutionary strategy by which workers can identify conspecific colonies, evade detection by guards, and avoid competition with genetic relatives.

Social environment affects sensory gene expression in ant larvae.

Social insects depend on communication to regulate social behaviour. This also applies to their larvae, which are commonly exposed to social interactions and can react to social stimulation. However, how social insect larvae sense their environment is not known. Using RNAseq, we characterized expression of sensory-related genes in larvae of the ant Formica fusca, upon exposure to two social environments: isolation without contact to other individuals, and stimulation via the presence of other developing individuals. Expression of key sensory-related genes was higher following social stimulation, and larvae expressed many of the same sensory-related genes as adult ants and larvae of other insects, including genes belonging to the major insect chemosensory gene families. Our study provides first insights into the molecular changes associated with social information perception in social insect larvae.

Vitellogenin and vitellogenin-like gene expression patterns in relation to caste and task in the ant Formica fusca

Social insect colonies are characterized by division of labour, and extensive morphological, physiological and behavioural differences between queens and workers. The storage protein vitellogenin (Vg) affects multiple aspects of social insect life histories, and has been suggested as a key player for caste differentiation and maintenance. Recently, three genes homologous to Vg have been described in the ant Formica exsecta. Their role is currently unclear but their structural variation suggests variable functions. We examined the expression patterns of the conventional Vg and the three Vg-like genes using qRT-PCR in the common black ant Formica fusca between queens and workers, between nurse and foragers workers, and across social contexts (queenless vs. queenright nests), and sampling time. As expected, we found a significant queen caste and nurse task-related increase for the conventional Vg, while Vg-like-C displayed a consistent forager-biased expression pattern. Task (forager vs. nurse) was the only factor that explained expression variation among workers in any of the studied genes. The removal of the queen did not affect expression, although the proportion of fertile nurses increased in queenless nests. The observed expression biases suggest that in Formica fusca, the ancestral duplication has led to alternative social functions for Vg-like genes across castes and tasks. To get a broader picture of the role of gene duplications in social evolution and the roles of Vg-like genes in caste differentiation and maintenance, how these genes achieve these roles at a molecular level need to be investigated further.

Induced immune responses in Formica fusca (Hymenoptera: Formicidae)

Parental immune experience can enhance offspring defence mechanisms towards prevalent pathogens in the surrounding environment. This process of inherited resistance from one generation to another is known as trans-generational immune priming (TGIP) in invertebrates. In sedentary and dense insect societies, such as ant colonies, TGIP can influence colony survival and fitness upon pathogen outbreaks. However, TGIP appears to depend on species and environmental stressors and therefore can vary in intensity, as well as in the molecular mechanisms leading to resistance. Here, we stimulated the immune system of queens of the ant Formica fusca (Linnaeus, 1758) by wounding or injecting dead conidia of the entomopathogenic fungus Beauveria bassiana (Bals.-Criv.) Vuill. (1912). The offspring were subsequently infected with B. bassiana , and the effect of this priming on survival was evaluated. Furthermore, we investigated whether immune challenge of the mother queen induces changes in the expression of immunity-related genes in queens themselves and their brood. We combined this information with measurements of offspring size and number. Larvae produced by untreated queens had a significantly higher mortality after infection with B. bassiana , whereas those produced by immune-primed queens survived no worse than unexposed ones. Adult worker offspring from sham-control mothers showed a protective effect of queen treatment, consistent with transgenerational immune priming. Thus, the effects of queen priming appear to manifest themselves slightly differently in larval and adult offspring. No differences were detected in offspring number or size, but immune gene expression levels showed changes, both in queens and their offspring. Open access, licensed under CC BY 4.0. © 2018 The Author(s). Pages: 53-66 DOI: 10.25849/myrmecol.news_028:053 Volume: 28 Year: 2018 Journal: Myrmecol. New

De novo transcriptome assembly and its annotation for the black ant Formica fusca at the larval stage

Communication and nutrition are major drivers of fitness in ants. While communication is paramount to colony cohesion, nutrition is decisive in regulating reproductive division of labor among colony members. However, neither of these has been studied from a molecular perspective in developing individuals. Here, we report the availability of the first transcriptome resources for larvae of the ant Formica fusca, a species with excellent discrimination abilities and thus the potential to become a model system for studying molecular mechanisms of communication. We generated a comprehensive, high-coverage RNA-seq data set using Illumina RNA-seq technology by sequencing 24 individual 1st - 2nd instar larvae collected from four experimental groups (6 samples per treatment, 49 million mean reads per sample, coverage between 194–253×). A total of 24,765 unigenes were generated using a combination of genome-guided and de novo transcriptome assembly. A comprehensive assembly pipeline and annotation lists are provided. This dataset adds valuable transcriptomic resources for further study of developmental gene expression, transcriptional regulation and functional gene activity in ant larvae.

Individual behavioural type and group performance in Formica fusca ants

The link between individual and group-level behaviour may help understanding cooperation and division of labour in social animals. Despite the recent surge of studies, especially in social insects, the way individual differences translate into group performance remains debated. One hypothesis is that groups may simply differ in the average personality of their members and this would translate into inter-group differences in collective behaviour. We tested the hypothesis of a linear relationship between individual and group phenotype in the ant Formica fusca by using same-age groups of workers after measuring an individual behavioural trait. Individual exploratory activity in an open-field arena was significantly repeatable. Based on this trait, groups were composed, each consisting of 6 individuals with similar exploration tendency housed with 3 cocoons and a refuge. Individual exploratory activity was associated with the performance in cocoon recovery at the group level: groups composed of high exploratory individuals started transporting displaced cocoons significantly earlier and transported more cocoons into the refuge than groups with low exploratory workers. When in a group, more exploratory animals showed significantly more returns to the refuge than less exploratory ones and tended to transport more cocoons. These results show a direct linear link between individual and collective behaviour, suggesting that colony personality reflects the average personality of workers involved in a given task.

Weak population structure in the ant Formica fusca.

Dispersal is a fundamental trait of a species’ biology. High dispersal results in weakly structured or even panmictic populations over large areas, whereas weak dispersal enables population differentiation and strong spatial structuring. We report on the genetic population structure in the polygyne ant Formica fusca and the relative contribution of the dispersing males and females to this. We sampled 12 localities across a ∼35 km2 study area in Finland and generated mitochondrial DNA (mtDNA) haplotype data and microsatellite data. First, we assessed queen dispersal by estimating population differentiation from mtDNA haplotype data. Second, we analysed nuclear DNA microsatellite data to determine overall population genetic substructure in the study area with principal components analysis, Bayesian clustering, hierarchical F statistics and testing for evidence of isolation-by-distance. Third, we directly compared genetic differentiation estimates from maternally inherited mtDNA and bi-parentally inherited DNA microsatellites to test for sex-bias in dispersal. Our results showed no significant spatial structure or isolation by distance in neither mtDNA nor DNA microsatellite data, suggesting high dispersal of both sexes across the study area. However, mitochondrial differentiation was weaker (Fst-mt = 0.0047) than nuclear differentiation (Fst-nuc = 0.027), which translates into a sixfold larger female migration rate compared to that of males. We conclude that the weak population substructure reflects high dispersal in both sexes, and it is consistent with F. fusca as a pioneer species exploiting unstable habitats in successional boreal forests.

Recognition of nestmate eggs in the ant Formica fusca is based on queen derived cues

Abstract Inclusive fitness benefits depend on recognizing the right individuals to interact with. Social insect nests protect themselves from non-kin intruders through nestmate recognition based on chemical cues. The recognition cues on adult individuals are from a mixture of genetic and environmental sources, but the ontogeny and use of recognition cues on eggs has not been previously assessed. We studied recognition by workers of eggs that were either nestmates or non-nestmates. and the ontogeny of recognition cues on eggs in the ant Formica fusca, a species with precise egg recognition abilities. Workers were able to discriminate among freshly laid eggs with no nest derived cues on them, and the egg surface chemicals varied among nests in these eggs, suggesting that queen derived cues are used in nestmate recognition. The results are discussed in the light of their implications on deceptive social parasite strategies and within colony conflicts.

Cuticular Chemistry of Males and Females in the Ant Formica fusca

Communication between organisms involves visual, auditory, and olfactory pathways. In solitary insects, chemical recognition cues are influenced mainly by selection regimes related to species recognition and sexual selection. In social insects, chemical recognition cues have an additional role in mediating recognition of society members and, thereby, allowing kin selection to operate. Here, we examined whether cuticular hydrocarbon profiles are sex-specific and whether males and young queens of the ant Formica fusca have colony-specific profiles. We also investigated whether there is a relationship between genetic relatedness and chemical diversity within colonies. We demonstrated that female and male sexuals do not have unique sex-specific compounds, but that there are quantitative chemical differences between the sexes. Out of the 51 cuticular hydrocarbon compounds identified, 10 showed a significant quantitative difference between males and females. We also showed that both males and females have a significant colony-specific component in their profiles. Finally, we found a negative correlation between within-colony relatedness and within-colony chemical diversity of branched, but not linear compounds. This suggests that colonies with multiple matri- or patrilines also have a significantly greater chemical diversity.

Genetic diversity, colony chemical phenotype, and nest mate recognition in the ant Formica fusca

Recognition of relatives is often crucial for adaptive social behavior, but availability of recognition cues may limit adaptation. Social insect workers direct altruism toward relatives through nest mate recognition. We studied whether genetic diversity increases nest mate recognition cue diversity and weakens nest mate recognition behavior in the ant Formica fusca that has both multiple and single queen societies in sympatric populations. Despite larger genetic diversity in multiple queen than single queen colonies, we found no differences in chemical recognition cue diversity or aggression toward non-nest mates in behavioral bioassays. The results suggest that the relationship between individual genotypes and the colony chemical phenotype is a complex interaction of genes and the environment, including the social setting of the colony and that increase in genetic diversity does not increase cue diversity to levels that would cause informational constraints on recognition behavior in the species. Key words: chemical diversity, Formica, genetic diversity, kin selection, nest mate recognition, social insects. [Behav Ecol]

Are you my mother? Kin recognition in the ant Formica fusca

In social insects, workers trade personal reproduction for indirect fitness returns from helping their mother rear collateral kin. Colony membership is generally used as a proxy for kin discrimination, but the question remains whether recognition allows workers to discriminate between kin and nonkin regardless of colony affiliation. We investigated whether workers of the ant Formica fusca can identify their mother when fostered with their mother, their sisters, a hetero-colonial queen or hetero-colonial workers. We found that workers always displayed less aggression towards both their mother and their foster queen, as compared to an unfamiliar hetero-colonial queen. In support of this finding, workers maintain their colony hydrocarbon profile regardless of foster regime, yet show modifications when exposed to different environments. This indicates that recognition entails environmental and genetic components, which allow both discrimination of kin in the absence of prior contact and learning of recognition cues based on group membership.

Genetic population structure, queen supersedure and social polymorphism in a social Hymenoptera

In social insects, the emergence of multiple queening is linked to changes in a suite of traits such as the reproductive life span of queens, mating patterns and population structure. We investigated queen turnover, colony longevity, spatial distribution patterns and genetic differentiation in a population of the socially polymorphic ant Formica fusca. Genetic differentiation between the social forms was absent, and mating patterns were similar in the two forms. The spatial distribution of single- and multi-queen colonies indicated an absence of colony reproduction by budding in both colony types. However, the rate of queen supersedure was high in multi-queen colonies and absent in single-queen ones. The social structure of colonies remained stable across years, but colony mortality did not differ between the two social forms. These results imply that differences between social types may appear and persist also in sympatry, and that these differences may occur in some traits, but not others, despite the presence of homogenizing gene flow.

Multiple breeders, breeder shifts and inclusive fitness returns in an ant

In social insects, colonies may contain multiple reproductively active queens. This leads to potential conflicts over the apportionment of brood maternity, especially with respect to the production of reproductive offspring. We investigated reproductive partitioning in offspring females (gynes) and workers in the ant Formica fusca, and combined this information with data on the genetic returns gained by workers. Our results provide the first evidence that differential reproductive partitioning among breeders can enhance the inclusive fitness returns for sterile individuals that tend non-descendant offspring. Two aspects of reproductive partitioning contribute to this outcome. First, significantly fewer mother queens contribute to gyne (new reproductive females) than to worker brood, such that relatedness increases from worker to gyne brood. Second, and more importantly, adult workers were significantly more related to the reproductive brood raised by the colony, than to the contemporary worker brood. Thus, the observed breeder shift leads to genetic benefits for the adult workers that tend the brood. Our results also have repercussions for genetic population analyses. Given the observed pattern of reproductive partitioning, estimates of effective population size based on worker and gyne samples are not interchangeable.

Worker policing and nest mate recognition in the ant Formica fusca

A conflict over male production arises in social insects where workers are able to lay unfertilized male eggs. This happens because each female (queen or worker) is most closely related to her own sons and is thus predicted to reproduce. The conflict is modulated by worker policing where workers prevent each other from reproducing by aggression or egg cannibalism. In this study, we show that in the ant Formica fusca, worker policing occurs by egg cannibalism rather than by overt aggression among workers. Furthermore, we show that, contrary to bees, wasps and other ant species, egg discrimination in F. fusca is not based only on a universal queen signature chemical and that nest mate recognition of eggs occurs.

Worker reproduction in the antFormica fusca

We studied the kin conflict over male parentage in the ant Formica fusca. The conflict arises because each worker and queen is most related to her own sons and is thus predicted to lay eggs. Microsatellite analysis of eggs revealed that workers laid eggs in more than half the queenright experimental nests. Nevertheless, almost exclusively diploid offspring were reared in the presence of a queen. This also occurred when worker-laid haploid male eggs were experimentally introduced in to the nests. Because our experimental setup allowed us to exclude the possibility of queen policing, we conclude that worker laid eggs are removed by other workers, either as a response to their parentage or gender. Our results suggest that worker reproduction in F. fusca is ultimately an interplay of conflicts over male parentage and sex allocation and that both worker and self policing have roles as proximate mechanisms of resolution.

Reproductive sharing among queens in the ant Formica fusca

Reproductive sharing among cobreeders, in which reproductive shares may vary from equal contribution (low reproductive skew) to reproductive dominance by one individual (high reproductive skew), is a fundamental feature of animal societies. Recent theoretical work, the reproductive skew models, has focused on factors affecting the degree to which reproduction is skewed within a society. We used the parameters provided by skew models as a guideline to study determinants of reproductive sharing in polygyne ants. As a model system we used two-queen laboratory colonies of the ant Formica fusca in which the reproductive shares of each queen was assessed from offspring by using allozymes and DNA microsatellites. We tested how the different variables included in reproductive skew models (queen-queen relatedness, potential fighting ability, productivity, and worker relatedness reflected by queen number in the colony of origin) affect reproductive sharing among queens. The results showed that the relatedness among queens explained 26% of the variation in reproductive skew. The size difference between queens (reflecting potential fighting ability), colony productivity, and worker relatedness did not have an effect on reproductive partitioning among cobreeders. To our knowledge, this is the first study to test for the effects of various determinants of skew in an experimental setting. Copyright 2003.

Sociobiology: Worker nepotism among polygynous ants.

Insect societies are a prime example of extreme cooperation, but their social life also entails the pursuit of selfish interests by society members1. Here we show that workers of the ant Formica fusca favour their own close kin when rearing eggs and larvae in colonies that are derived from several queens. This nepotistic behaviour indicates that ant workers are able not only to detect kin relationships, but also to pursue their selfish genetic interests if the costs to their colony are not prohibitive.

Proximate Determinants of Reproductive Skew in Polygyne Colonies of the AntFormica fusca

AbstractUnderstanding the determinants of reproductive skew (the partitioning of reproduction among co‐breeding individuals) is one of the major questions in social evolution. In ants, multiple‐queen nests are common and reproductive skew among queens has been shown to vary tremendously both within and between species. Proximate determinants of skew may be related to both queen and worker behaviour. Queens may attempt to change their reproductive share through dominance interactions, egg eating and by changing individual fecundity. Conversely, workers are in a position to regulate the reproductive output of queens when rearing the brood. This paper investigates queen behaviour at the onset of egg laying and the effect of queen fecundity and worker behaviour on brood development and reproductive shares of multiple queens in the antFormica fusca. The study was conducted in two‐queen laboratory colonies where the queens produced only worker offspring. The results show that in this species reproductive apportionment among queens is not based on dominance behaviour and aggression, but rather on differences in queen fecundity. We also show that, although the queen fecundity at the onset of brood rearing is a good indicator of her final reproductive output, changes in brood composition occur during brood development. Our results highlight the importance of queen fecundity as a major determinant of her reproductive success. They furthermore suggest that in highly derived polygyne species, such as theFormicaants, direct interactions as a means for gaining reproductive dominance have lost their importance.