Wood Ant Formica Research Articles

Do They Know What They Are Doing? Cognitive Aspects of Rescue Behaviour Directed by Workers of the Red Wood Ant Formica polyctena to Nestmate Victims Entrapped in Artificial Snares.

Ant rescue behaviour belongs to the most interesting subcategories of prosocial and altruistic behaviour encountered in the animal world. Several studies suggested that ants are able to identify what exactly restrains the movements of another individual and to direct their rescue behaviour precisely to that object. To shed more light on the question of how precise the identification of the source of restraint of another ant is, we investigated rescue behaviour of red wood ant Formica polyctena workers, using a new version of an artificial snare bioassay in which a nestmate victim bore two wire loops on its body, one (acting as a snare) placed on its petiole and an additional one on its leg. The tested ants did not preferentially direct their rescue behaviour towards the snare. Moreover, the overall strategy adopted by the most active rescuers was not limited to precisely targeted rescue attempts directed towards the snare, but consisted of frequent switching between various subcategories of rescue behaviour. These findings highlight the importance of precise identification of cognitive processes and overall behavioural strategies for better understanding of causal factors underlying animal helping behaviour in light of new facts discovered by testing of various successive research hypotheses.

Comparative Analysis of Facial Coloration between Introduced and Source Populations of the Red Wood Ant Formica paralugubris.

The variation in the typical black-reddish color of red wood ants (Formica rufa group) has been recently suggested as a good indicator of habitat quality, being dependent on environmental conditions. However, the relative contribution of external factors and heritability in shaping this trait is poorly investigated. In this study, we compared the facial coloration of workers from four introduced populations of Formica paralugubris with those of the two Alpine populations from which they had been taken. We used a Relative Warp Analysis to describe the variations in the shape of this trait. We expected each introduced population to be more similar to its population of origin if the color pattern was predominantly genetically determined. On the contrary, due to the considerable differences in habitat type and climate between the Alps and the Apennines, we expected to observe differences between the introduced population and their origin population if the coloration was mostly environmentally determined. With one exception that we discuss, the results showed that ants from the two source populations had different phenotypes, and that the introduced populations had a shape similar to the population of origin, suggesting a stable genetic background. Surprisingly, the habitat type seems to have a less clear effect, even if within-population differences suggest the influence of very localized environmental factors. Finally, we found that the facial coloration shape is affected by the ant’s size, a result in line with previous studies.

The Effect of the Introduced Red Wood Ant Formica paralugubris on the Frequency of Ant Nests and First Plesiobiotic Association between Myrmica ruginodis (Nylander, 1846) and Lasius flavus (Fabricius, 1782)

Red wood ants are ecologically dominant species that affect the composition of the invertebrate community. In the past century, one of these species, Formica paralugubris (Seifert 1996), was introduced outside its native area (the Alps), in the Apennines (Italy). Here we investigated the effect of the presence of an introduced population of F. paralugubris on the frequency of nests of other ant species in Abetone, Central Apennines. Ant nests were found only in the area unoccupied by F. paralugubris. In this area, we also found one pleometrotic association between queens of Lasius flavus (Fabricius, 1782) and two plesiobiotic associations, one between Formica fusca (Linnaeus, 1758) and L. flavus and the other one between Myrmica ruginodis (Nylander, 1846) and L. flavus. This latter is the first plesiobiotic relationship reported between these two species, the second reported for M. ruginodis and the 10th for L. flavus. Additionally, this is the first plesiobiotic association reported for the Mediterranean region and Southern Europe.

The Effect of the Introduced Red Wood Ant Formica paralugubris on the Frequency of Ant Nests and First Plesiobiotic Association between Myrmica ruginodis (Nylander, 1846) and Lasius flavus (Fabricius, 1782)

Red wood ants are ecologically dominant species that affect the composition of the invertebrate community. In the past century, one of these species, Formica paralugubris (Seifert 1996), was introduced outside its native area (the Alps), in the Apennines (Italy). Here we investigated the effect of the presence of an introduced population of F. paralugubris on the frequency of nests of other ant species in Abetone, Central Apennines. Ant nests were found only in the area unoccupied by F. paralugubris. In this area, we also found one pleometrotic association between queens of Lasius flavus (Fabricius, 1782) and two plesiobiotic associations, one between Formica fusca (Linnaeus, 1758) and L. flavus and the other one between Myrmica ruginodis (Nylander, 1846) and L. flavus. This latter is the first plesiobiotic relationship reported between these two species, the second reported for M. ruginodis and the 10th for L. flavus. Additionally, this is the first plesiobiotic association reported for the Mediterranean region and Southern Europe.

Assessing Molecular Diversity in Native and Introduced Populations of Red Wood Ant Formica paralugubris.

The Formica rufa group comprises several ant species which are collectively referred to as "red wood ants" and play key roles in boreal forest ecosystems, where they are ecologically dominant and greatly influence habitat dynamics. Owing to their intense predatory activity, some of these species are used as biocontrol agents against several forest insect pests and for this aim in Italy, nearly 6000 ant nests were introduced from their native areas in the Alps to several Appeninic sites during the last century. In this work, we assessed and compared the genetic variability and structure of native and introduced populations of F. paralugubris, thus evaluating the extent of genetic drift that may have occurred since the time of introduction, using amplified fragment length polymorphism (AFLP) markers. PCR amplification with a fam_EcoRI-TAC/MseI-ATG primers combination produced a total of 147 scorable bands, with 17 identified as outlier loci. The genetic variation was higher in the introduced population compared to the native ones that, on the other hand, showed a higher diversity between nests. AMOVA results clearly pointed out that the overall genetic structure was dominated by among-worker variation, considering all populations, the Alpine vs. Apennine groups and the comparison among native and related introduced populations (all ranging between 77.84% and 79.84%). Genetic analyses unveiled the existence of six main different groups that do not entirely mirror their geographic subdivision, pointing towards a wide admixture between populations, but, at the same time, rapid diversification of some Apennine populations. Future studies based on high-throughput genomic methods are needed to obtain a thorough understanding of the effects of environmental pressure on the genetic structure and mating system of these populations.

Intraspecific Relationships and Nest Mound Shape Are Affected by Habitat Features in Introduced Populations of the Red Wood Ant Formica paralugubris.

Simple SummaryRed wood ants (Formica rufa goup) are dominant ant species widespread in the Eurasian continent. These species have a strong ecological impact on the habitats they dwell in, being top-ranked predators. One of the most striking features of these ants is represented by the large nest mounds they build. In this study, we investigated how nest mound shape and colony organization of imported populations of Formica paralugubris varied in three different habitat types. We found that nest mounds differed in size, number and shape in the three habitats. In all the three sites, nests were connected by trails of workers, but the size of these nest-networks differed. We also investigated the pattern of intraspecific aggression among ants from different nests, and we showed that aggressiveness was higher within each population than between separate populations, a finding in line with a ‘nasty neighbor’ behavior.Ants belonging to the Formica rufa group build large nest mounds, which aid their survival during severe winters. We investigated whether different environmental features of the habitats affected the nest mound shape and the population structure. We assessed the shape of all the nest mounds and mapped inter-nest trails connecting mounds for three imported populations of Formica paralugubris in three forest habitats: fir-dominated, beech-dominated, and a mixture of fir and beech. Single-nest mounds were averagely smaller and flatter in the beech-dominated forest, probably because of lighter building materials. Nonetheless, by summing the volumes of all interconnected nests, the size was similar among all three sites. In fir- and beech-dominated forests, large nests were also central in the networks, suggesting a central place foraging model with these nests as reference. We finally performed aggression tests, and found that aggressiveness was significantly higher among nests belonging to the same population than between populations. The results highlight the plasticity of the species to adapt nest and colony structure to different environments. Additionally, it appears that none of these populations is unicolonial, as observed in various alpine sites, there and the observed patterns of aggression are coherent with the ‘nasty neighbor’ effect.

Forest logging is associated with shape of males and gynes in the wood ant Formica aquilonia

Environmental stress can affect individual development and fitness in insects. Forest logging is a serious environmental stress for forest-specialist insects, such as the mound-building wood ant Formica aquilonia Yarrow 1955, which builds its nests into forests and is dependent on the nutrition provided by the forest habitat. We studied whether the logging causes such a strong environmental stress that it would be visible in disproportionate growth and shape of young winged wood ant gynes (‘queens’) and males. We measured head width, thorax width and wing length of 144 gynes and 100 males from 12 nests from six clear-cut areas, 11 nests from unlogged forest stands and 10 nests in clear-cut—forest margins. We observed disproportionate growth of different body parts and allometric growth of wings in both sexes in areas with different logging-induced disturbance. Gynes had larger heads in clear-cut areas and males had narrowest thoraxes in forest clear-cut edges. With an increasing thorax width, the wing length decreases steeper in clear-cuts than in other habitats in gynes and increases steeper in clear-cuts than other habitats in males. It seems that in a strongly disturbed clear-cut environment, the gynes invest the growth of head width. There seems to be a trade-off between the growth of the thorax and wings in gynes but not in males. The altered body shape may be adaptation to clearings caused by storm events of wild fires, but not forest management practices of modern era. Large-headed gynes may be better in a colony take-over, needed for the establishment of new colonies in early succession stage habitats in this temporally social parasitic species. Long-winged males may have better long-range flight ability, and they may thus have better fitness and change to disperse their genes onto new habitats.

CO2 biogeochemical investigation and microbial characterization of red wood ant mounds in a Southern Europe montane forest

Red wood ants are ecologically important species in Europe that form large colonies. Their nest mounds are characterized by stable microclimatic conditions, that are favourable to the development of rich invertebrate and microbial communities. Through their respiration processes, all these inhabitants contribute to the total gas emissions of the mounds. Quantifications of red wood ant mounds CO2 production are only available from Northern and Central Europe, and the Alps, where these ants are common. During the second half of the last century some species were transplanted from the Alps to southernmost sites, where they were not present, to be employed as biocontrol agents. No information on the contribution of these low-latitudes populations to the local forest CO2 production is available. The microbial communities living within red wood ant mounds are also poorly known. In this study, we investigated the CO2 gas emissions and the microbiome of the mounds of an introduced population of the red wood ant Formica paralugubris in a Southern Europe montane forest. We found that ant mounds produced more CO2 than the forest soil, and that their CO2 efflux as well as internal concentration were higher during summer than winter, with a lighter CO2 carbon isotopic signature in summer than winter, likely due to an increased ant activity. Moreover, the top part of the mound was characterised by higher CO2 efflux and lower CO2 internal concentration compared to the bottom, probably due to its internal structure and conditions. The isotopic signature of the mound material was similar between summer and winter, suggesting a metabolic similarity of the microbial communities. Also, we estimated the ants’ relative contribution to the total mound CO2 production to be 83%, whereas the microbiota CO2 contribution was estimated at 17%. Finally, the mound microbiome composition varied between summer and winter, though no seasonal difference in the diversity indexes or β-diversity was found. Our results demonstrate the impacts of the introduced red wood ants on the carbon dynamics of the recipient ecosystem.

Terpenes and fungal biomass in the nest mounds of Formica aquilonia wood ants

Nest mounds of wood ants of the Formica rufa group are built using conifer needles, small branches, other plant materials and soil. Conifer needles contain several mono and sesquiterpenes. Thus, wood ant nests may act as terpene hotspots in conifer forest soils. Some of the terpenes show antifungal activity and may thus cause small-scale heterogeneity in fungal biomass. We compared terpene concentrations and fungal biomass (ergosterol concentration) of nest material of the wood ant Formica aquilonia from the top, core and basement of 14 nest mounds in eastern Finland. Overall, α-pinene, camphene, sabinene, β-pinene, myrcene, limonene, camphor and longifolene were detected the most and were commonly present in all the studied layers. We found that terpene concentrations differed between the sampled nest material layers, being generally highest in the core of the nest and lowest at the basement of the nest. There was no association between the terpene concentration and material moisture. Fungal biomass was highest in the top layer, intermediate in core and lowest in basement; however, it was not negatively associated with terpene concentrations. Fungal biomass in nest mounds was positively associated with moisture and alkalinity. Nest mounds of Formica rufa group wood ants are complex structures with different chemical and microbial properties among its layers.

A comparative study of the fauna associated with nest mounds of native and introduced populations of the red wood ant Formica paralugubris

In the second half of the twentieth century, many red wood ant populations were transferred from the Alps to the Apennines as biological control agents. Since the introduction involved the relocation of entire nest mounds, it is presumable that the associated fauna was also relocated. While the introduction of these ants has raised several concerns about their ecological impact, there has been no attempt to investigate the introduction of other nest-associated species. In this study, we collected samples of soil and nest material from three populations of the red wood ant Formica paralugubris, one Alpine native and two imported into the Apennines. We aimed to confirm that nest mounds are hotspots for soil fauna, detect the occurrence of new myrmecophilous species, and compare the nest-associated fauna among sites, to test the hypothesis of mass species relocation. We focused our analyses mainly on two taxa, springtails and oribatid mites, two highly representative groups of the mesofauna inhabiting nest mounds. The results showed higher richness and diversity in nests than soil for oribatids but not for springtails. We found 17 myrmecophilous oribatid species, but only two springtail species. Finally, native and imported sites shared only a few oribatid and springtail species, suggesting that massive relocation did not occur with nest transplants or it was likely limited. Additionally, we found some species never before collected in Italy.

Competitive pressure by territorials promotes the utilization of unusual food source by subordinate ants in temperate European woodlands

In the presence of territorial dominant ant species, lower-ranked ants change their foraging strategy by shifting their activity in time and space and by switching to suboptimal food sources. Although important for many insect species, pollen is a rarely used food source by ants. Occasional pollen consumption was demonstrated only in a handful of ant species that are mainly lower-ranked in the competitive hierarchy. We hypothesized that the consumption of pollen by subordinate ants might be enhanced by competitive pressure exerted by dominant species. In the frame of the current study, we assessed the food preference of ant species within a supercolony of the territorial wood ant Formica polyctena by offering them pollen and animal protein baits. Subordinate ant species, especially Myrmica ants, exploited both pollen and animal protein, while F. polyctena foraged only for animal protein. The Myrmica ants exploited pollen sources more intensively on plots where F. polyctena dominated animal protein baits, thus interspecific competition was stronger. As suggested by our study, under suboptimal conditions, where food availability is hindered, e.g. by the presence of superior competitors, pollen can, indeed, be used as a food source by subordinate ants.

Living on the Edge: Changes in the Foraging Strategy of a Territorial Ant Species Occurring with a Rival Supercolony \u2013 a Case Study

Territorial strategy in animals is characterized by the monopoly of resources inside a protected area, the territory. The presence of territorial species considerably alters the behavior of co-occurring submissives, as it is known in several submissive ant species living on the territories of red wood ants in temperate regions. On the other hand, as a rule, territorial species cannot share the same territory and usually exclude each other. However, this ‘rule’ is inferred from the almost complete lack of data on the coexistence of rival territorials, and not from observations regarding the effective exclusion or behavioral inhibition of one territorial species by other. In the frame of this study, we investigated the foraging strategy of the territorial red wood ant Formica pratensis that occurred inside a large polydomous system of another territorial ant species, F. exsecta. Formica pratensis colonies outside the supercolony served as control. Within the F. exsecta supercolony, F. pratensis showed the characteristics of a subordinate species characterized by low discovery and exploitation success of artificial food sources. On the contrary, control colonies outside the polydomous system clearly behaved like typical territorials as they successfully monopolized the majority of the baits. In addition, submissive species were more successful around the F. pratensis nests within the supercolony, than outside of it. As suggested by our results, territorial species could co-occur with other territorials given certain plasticity in their behavior and small colony size. Nevertheless, exclusion also happens as proven by our field observations.

Body Size of Wood Ant Workers Affects their Work Division

The division of functions among ant workers and their mutual cooperation is one of the reasons for ants’ success. The activities that workers perform in the nest can be divided by age or morphology. We studied the body size of workers of the wood ant Formica polyctena as a function of their activity. Our results show that workers exploiting protein baits were larger than workers attracted to carbohydrate baits. The biggest of all were workers located at the upper parts of the nest, which shows the importance of nest defense and maintenance. It also points that the distribution of functions does not have to be given only by workers age. Work division based on several mechanisms may be advantageous for colony functioning.

Ants trapped for years in an old bunker; survival by cannibalism and eventual escape

Successful evacuation of a peculiar ‘colony’ of the wood ant Formica polyctena Först., for years trapped within an old bunker previously used for storing nuclear weapons (see Czechowski et al. 2016), is reported. Using an experimentally installed boardwalk, the imprisoned ants managed to get through the ventilation pipe to their maternal nest on the top of the bunker. In our previous report, we left open the question of how the ‘colony’ could survive seemingly without food. Here we show that the ‘colony’ in the bunker survived and grew thanks to an influx of workers from the source nest above the bunker and mass consumption of corpses of the imprisoned nestmates.

The impacts of forest clear felling on the oribatid mite fauna inhabiting Formica aquilonia nest mounds

The ubiquitous soil-dwelling oribatid mites have long been investigated as potential bioindicator agents for environmental changes, such as those resulting from forest management. However, the oribatid community composition often lacks detectable changes, mainly due to the difficulties of finding homogenous microhabitats from the landscape for investigation purposes. Red wood ant nest mounds in boreal forests, however, form uniform, moisture, and temperature regulated, microhabitats of which are also inhabited by a rather characteristic oribatid fauna. With this standardized microhabitat—the red wood ant Formica aquilonia nest mounds in Finland—we studied the impacts of forest clear felling on the oribatid mites inhabiting those nests. We compared the oribatid assemblages of altogether 41 ant mounds, located in three separate clear fell areas and in adjacent mature coniferous forest. A total of 16,499 specimens, representing 67 oribatid species, were observed. The ten most abundant species represented 90.9% of the total abundance. The oribatid species richness was significantly lower in clear fell mounds and was positively related to the surface moisture content of the mounds. Oribatid abundance or community composition did not markedly differ between clear fell and forest mounds. Despite the attempt to standardize the microhabitat of oribatid fauna, the use of oribatid mites as biological indicators of harvesting disturbance was limited with this study design due the lack of changes in community composition. Then again, the oribatid species richness of this characteristic microhabitat may provide useable quality measures about the harms of forest clear felling.

The first draft genomes of the ant Formica exsecta, and its Wolbachia endosymbiont reveal extensive gene transfer from endosymbiont to host

BackgroundAdapting to changes in the environment is the foundation of species survival, and is usually thought to be a gradual process. However, transposable elements (TEs), epigenetic modifications, and/or genetic material acquired from other organisms by means of horizontal gene transfer (HGTs), can also lead to novel adaptive traits. Social insects form dense societies, which attract and maintain extra- and intracellular accessory inhabitants, which may facilitate gene transfer between species. The wood ant Formica exsecta (Formicidae; Hymenoptera), is a common ant species throughout the Palearctic region. The species is a well-established model for studies of ecological characteristics and evolutionary conflict.ResultsIn this study, we sequenced and assembled draft genomes for F. exsecta and its endosymbiont Wolbachia. The F. exsecta draft genome is 277.7 Mb long; we identify 13,767 protein coding genes, for which we provide gene ontology and protein domain annotations. This is also the first report of a Wolbachia genome from ants, and provides insights into the phylogenetic position of this endosymbiont. We also identified multiple horizontal gene transfer events (HGTs) from Wolbachia to F. exsecta. Some of these HGTs have also occurred in parallel in multiple other insect genomes, highlighting the extent of HGTs in eukaryotes.ConclusionWe present the first draft genome of ant F. exsecta, and its endosymbiont Wolbachia (wFex), and show considerable rates of gene transfer from the symbiont to the host. We expect that especially the F. exsecta genome will be valuable resource in further exploration of the molecular basis of the evolution of social organization.

The Preference of Some Myrmecochorous Plants of Forest Stands by Red Wood Ant (Formica rufa L.)—Experiment on Seeds with Elaiosomes

The paper is aimed in ants’ preference among seeds with elaiosomes. Seeds with elaiosomes of plants Asarum europaeum, Viola arvensis, Symphytum officinale, Chelidonium majus, and Ficaria verna were offered to red wood ants (Formica rufa) near two observed anthills. Monitored characteristics were the number of seeds collected by ants depending on the plant species, the presence of elaiosome, the orientation and the distance from anthill. The obtained data were processed by factorial ANOVA including PCA analysis. Seeds without elaiosomes were less preferred than the seeds with elaiosomes at statistical level. This experiment confirmed that elaiosomes attached to the seeds had significantly influenced transport of seeds by red wood ants. On the contrary, the influence of orientations (north–south) between seeds and the anthill were not proved. A distance of seeds from the anthill may play a certain role, but the results were not significant. Data indicated that ants had preferred seeds with elaiosomes of two plant species (A. europaeum and V. arvensis) from five species offered. The authors recommend to store seeds in a freezer before a similar experiment, because the elaisomes from different plant species will be fresh.

Effects of genetic relatedness, spatial distance, and context on intraspecifi c aggression in the red wood ant Formica pratensis (Hymenoptera: Formicidae)

In the present study, we tested the level of aggression of monodomous and polydomous colonies of the wood ant Formica pratensis (Hymenoptera: Formicidae) with a behavioral assay in nature and laboratory conditions to see if the ants from neighboring colonies are more tolerant or more aggressive to each other than those from greater distances. We also tested how context (nature and laboratory conditions) affected aggression. Our results showed that the monodomous colonies were highly aggressive to all neighboring or nonneighboring conspecifics in nature irrespective of the spatial distance. The polydomous colony showed no aggression towards neighboring conspecifics but the level of aggression increased with increasing spatial distance between the colonies. The level of aggression of tested colonies in laboratory conditions was significantly low, irrespective of whether they were monodomous or polydomous, indicating that aggression is context dependent. A DNA barcoding technique based on mitochondrial COI gene sequencing was applied to determine the genetic relatedness between the colonies. The results of the genetic analysis, in combination with results of behavioral assays, revealed that aggression behavior of the polydomous colony was affected by both the genetic relatedness and the spatial distance between the colonies while there was no clear separation of effects of genetic relatedness and spatial distance on aggression in the monodomous colonies.

Distribution of oribatid mites is moisture-related within red wood ant Formica polyctena nest mounds

Abundant and species rich oribatid mite faunas live as ant associates in red wood ant nest mounds. Distribution of oribatids (as of many other groups) within mounds is unknown, yet there is an indication that many oribatid species favor moist microclimates. The ant mounds typically have a relatively dry core covered by a moister surface layer, but the moisture variance of mound parts has rarely been verified by measurements. We measured the moisture variance of nine Formica polyctena mounds (surface, rim, core) located in three different forest types in Finland. The moisture may affect the mounds’ temperature maintenance, and we further investigated this relationship. We also hypothesised that the abundance, species richness and community composition of oribatids may be related to the moisture, and investigated these differences between the mound parts and forest types. The moisture varied significantly between the mound parts; moisture was highest in surface followed by rim and core. The surface moisture had a positive relationship with nest core temperature. A total of 18,614 oribatid mites belonging to 93 species and one higher taxon were recorded. The oribatids occurred predominantly on the surface (77.9%), followed by the rim (16.6%) and the core (5.5%) of the ant nest mounds. The abundance and species richness of oribatids was positively related to moisture and differed significantly between the mound parts and forest types. The mound volume had no effect on moisture, temperature or mite community composition. The results indicate that the moisture harbored in the surface layer of ant nest mounds is a vital factor for maintaining rich oribatid mite communities within mounds.

Not simply red: Colouration of red wood ant Formica rufa (Hymenoptera: Formicidae) is polymorphic, modular and size-dependent

Insects produce pigment and structural colours mainly for camouflage, signaling, physical protection or temperature regulation, and colour patterns can provide information about individual quality. Although the evolutionary function and nature of the variability in colouration are well known for many invertebrate taxa, there is little information on this topic for ants. We studied individual variation in the melanin-based colour traits of workers of the red wood ant, Formica rufa (Hymenoptera: Formicidae), from 20 colonies in Southern Finland and revealed the type of colouration in this species. First, using the threshold approach we distinguished between continuous and discrete variations. Furthermore, the analyses affirmed nine discrete morphs in terms of the colouration on the head and eight on the pronotum, while only continuous variation were found on the other body parts. Measuring the size of a particular colour pattern, the intensity of colour expression (degree of melanization) and statistical analyses allowed an assessment of the intra-individual variation in both discrete and continuous patterns. The results revealed substantial modularity in the above mentioned colouration traits. In workers of F. rufa there were individuals with a dark head and light coloured thorax and vice a versa. Size of the dark pigment colour patterns exhibited less modularity than the degree of melanization. Finally, the interrelation between colouration traits and individual body size revealed their size-dependent origin. Small individuals had relatively larger areas of colour on the head and thorax than big individuals. These results are likely to facilitate further taxonomical and ecological studies on red wood ants, as they show it is possible to assess colouration traits in ants. However, more studies are needed on the function of polymorphism and modular colouration in this group of ants.