Wood Ants Research Articles

Impact of central complex lesions on innate and learnt visual navigation in ants

Wood ants are excellent navigators, using a combination of innate and learnt navigational strategies to travel between their nest and feeding sites. Visual navigation in ants has been studied extensively, however, we have little direct evidence for the underlying neural mechanisms. Here, we perform lateralized mechanical lesions in the central complex (CX) of wood ants, a midline structure known to allow an insect to keep track of the direction of sensory cues relative to its own orientation and to control movement. We lesioned two groups of ants and observed their behaviour in an arena with a large visual landmark present. The first group of ants were naïve and when intact such ants show a clear innate attraction to the conspicuous landmark. The second group of ants were trained to aim to a food location to the side of the landmark. The general heading of naïve ants towards a visual cue was not altered by the lesions, but the heading of ants trained to a landmark adjacent food position was affected. Thus, CX lesions had a specific impact on learnt visual guidance. We also observed that lateralised lesions altered the fine details of turning with lesioned ants spending less time turning to the side ipsilateral of the lesion. The results confirm the role of the CX in turn control and highlight its important role in the implementation of learnt behaviours that rely on information from other brain regions.

Bark Beetle Attacks Reduce Survival of Wood Ant Nests

Central European coniferous forests are facing the most significant bark beetle outbreak in history. The consequence is the creation of large clearings, which are associated with changes in environmental parameters. We studied how these changes affected the survival of wood ant nests. Specifically, we monitored the relationship between mound occurrence, tree coverage, and the coverage and height of vegetation on ant mounds. We found that wood ant nests were less likely to survive (39% decline) in bark beetle-affected forests than in unaffected forests. The Formica rufa species was more affected than F. polyctena. The bark beetle attack was significantly positively correlated with increases in understory vegetation cover and height. Several factors may explain the negative effects of bark beetles on wood ants. The removal of trees subsequently increased the amount of light on ground vegetation. The support of understory vegetation growth on ant mounds cast shadows on the nests and disrupted nest thermoregulation. Additionally, the bark beetle attack may have been associated with a loss of food resources for wood ants. Our results showed that early (small-scale) management intervention against the spread of bark beetles is the preferred method for maintaining the ant population in conifer forests prone to bark beetle attacks.

Predation risk estimated on live and artificial insect prey follows different patterns.

Models mimicking prey organisms are increasingly used in ecological studies, including testing fundamental ecological and evolutionary theories. The general consensus is that predation risk estimated on artificial models may not quantitatively correspond to predation pressure on live prey, but it still can be used in various comparisons. We tested whether the use of live and artificial prey reveals the same patterns of variation in predation risk. We exposed live prey (blowfly larvae and puparia) and plasticine models of blowfly puparia in two boreal forest sites, both openly and in ant- and bird-exclusion treatments, and we quantified attacks by both avian and invertebrate predators. Bird attack rates were always higher on live puparia than on their plasticine models, but the magnitude of this difference declined from 8.4-fold in early summer to 2-fold in mid- and late-summer. We attribute these changes to different responses to prey by experienced adult birds that dominate the bird communities in early summer versus explorative juvenile birds that are abundant later in the season. Invertebrate daily predation rates on maggots decreased from 56% in early summer to 28% in late summer, but invertebrate attacks on plasticine models showed no seasonal changes. Overall, invertebrate predation on maggots was 67-fold greater than their predation on models. Observations showed that wood ants did not attack plasticine models and did not leave on them any damage marks. Estimates based on artificial prey indicate a much greater role of bird predation than invertebrate predation, while estimates based on live prey suggest the opposite pattern. Thus, using live and artificial prey may lead to different conclusions about relative importance of different predator groups in a locality. Moreover, for both avian and invertebrate predators, predation risk based on artificial and live prey shows different seasonal changes and may potentially demonstrate different spatial patterns.

Differences Between the Allometric Rules Governing Two Formica lugubris Zetterstedt, 1838 (Hymenoptera: Formicidae) Ants Populations in Northern Greece

Red wood ant workers, despite being monomorphic, show a wide range in their body size even within the same colony. As overall size grows, some organs grow in a negative allometric relationship with the body. Genetic and environmental factors, like diet and temperature, are considered as key factors that determine differences in body and organ growth. We studied the allometric rules governing the organs in two different populations of red wood ants in two different environments in Northern Greece to identify similarities and differences. These rules, although linear for both populations and negative allometric for most of the traits measured, differ in the two study areas, while the shape of the head found to be governed by different allometric rules between the two populations. The whole shape and size of red wood ants varies greatly even within the same species and we cannot create allometric models for two different areas, even when they belong in the same species. The differences might be a result of differences in the genome and the environmental conditions between the two populations.

Changes in the Structure of Nest Complexes of the Red Wood Ants Formica rufa and F. polyctena (Hymenoptera, Formicidae) in Urban Forests

The aim of this work is to evaluate changes in the structure of nest complexes of two species of red wood ants: Formica rufa Linnaeus, 1761 and F. polyctena Foerster, 1850, in the city of Kyiv (Ukraine) using morpho[1]metric indicators, as well as to evaluate their dynamics. In 2014–2022, nest complexes of two species of red wood ants (Formica rufa, F. polyctena) were observed in urbanised forests within the city of Kyiv. A total of 472 F. rufa and 411 F. polyctena nests were studied in 7 nest complexes (4 — F. polyctena, 3 — F. rufa). The total volume of F. polyctena nests were found to decrease by 2020–2021, while the volume of F. rufa nests remained at the same level or even increased. This suggests that F. rufa is more resistant to changes in forestry conditions.

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.

Comparison of Geogases in Two Cenozoic Sedimentary Basins

We investigated fault gases (helium, radon, CO2) in two seismically active Cenozoic sedimentary basins: (a) Meinweg (in 2015), at a tectonically quiescent horst structure in the Lower Rhine Embayment; and (b) Bodanrück (in 2012; Lake of Constance), in the Molasse Basin and part of the seismically active Freiburg–Bonndorf–Bodensee Fault Zone (FBBFZ). Both study areas were selected because recent “GeoBio-Interactions” findings showed that red wood ants (RWA) are biological indicators of otherwise undetected degassing systems. We combined presence/absence data of RWA nests, their spatial pattern analysis (prototype lines), seismicity and known tectonic settings with soil gas analyses (a total of 817 samples) to unveil geochemical anomalies related to tectonic developments unknown so far. Currently, Meinweg can be considered “no ants land” due to the very low background-level of geogas concentrations. Thus, anomalies (Rn-CO2) weakly trending in NE-SW extension direction emerged. This could probably indicate the onset of (re)activation of the NE-SW-trending (Variscan) structures or the development of new fractures as an aftershock process of the 1992 Roermond earthquake that occurred about 15 km to the west. Results at Bodanrück (three RWA clusters and two RWA-free corridors) revealed degassing patterns in NW-SE and NNE-SSW directions in the clusters corresponding to re-activated and recent strike-slip fault systems. No gas anomalies were found in RWA-free corridors. The RWA nest distribution was shown to be a valuable tool for identifying areas of even actively degassing spotty anomalies caused by macro- and microscale brittle deformation masked by sediment cover.

Wood ants learn the magnetic direction of a route but express uncertainty because of competing directional cues.

ABSTRACTWood ants were trained indoors to follow a magnetically specified route that went from the centre of an arena to a drop of sucrose at the edge. The arena, placed in a white cylinder, was in the centre of a 3D coil system generating an inclined Earth-strength magnetic field in any horizontal direction. The specified direction was rotated between each trial. The ants’ knowledge of the route was tested in trials without food. Tests given early in the day, before any training, show that ants remember the magnetic route direction overnight. During the first 2 s of a test, ants mostly faced in the specified direction, but thereafter were often misdirected, with a tendency to face briefly in the opposite direction. Uncertainty about the correct path to take may stem in part from competing directional cues linked to the room. In addition to facing along the route, there is evidence that ants develop magnetically directed home and food vectors dependent upon path integration. A second experiment asked whether ants can use magnetic information contextually. In contrast to honeybees given a similar task, ants failed this test. Overall, we conclude that magnetic directional cues can be sufficient for route learning.

Innate visual attraction in wood ants is a hardwired behavior seen across different motivational and ecological contexts

Ants are expert navigators combining innate and learnt navigational strategies. Whereas we know that the ants’ feeding state segregates visual–navigational memories in ants navigating along a learnt route, it is an open question if the motivational state also affects the ants’ innate visual preferences. Wood ant foragers show an innate attraction to conspicuous visual cues. These foragers inhabit cluttered woodland habitat and feed on honeydew from aphids on trees. Hence, the attraction to ‘tree-like’ objects might be an ecologically relevant behavior that is tailored to the wood ants’ foraging ecology. Foragers from other ant species with different foraging ecologies show very different innate attractions. We investigated here the innate visual response of wood ant foragers with different motivational states, i.e., unfed or fed, as well as males that show no foraging activity. Our results show that ants from all three groups orient toward a prominent visual cue, i.e., this intrinsic visuomotor response is not context-dependent, but a hardwired behavior seen across different motivational and ecological contexts.

Dissecting the costs of a facultative symbiosis in an isopod living with ants.

The balance between costs and benefits is expected to drive associations between species. While these balances are well understood for strict associations, we have no insights to which extent they determine facultative associations between species. Here, we quantified the costs of living in a facultative association, by studying the effects of red wood ants on the facultatively associated isopod Porcellio scaber. Porcellio scaber frequently occurred in and near hostile red wood ant nests and might outnumber obligate nest associates. The facultative association involved different costs for the isopod. We found that the density of the isopod decreases near the nest with higher ant traffic. Individuals in and near the nest were smaller than individuals further away from the nest. Smaller individuals were also found at sites with higher ant traffic. A higher proportion of wounded individuals was found closer to the nest and with higher ant traffic. We recorded pregnant females and juveniles in the nest suggesting that the life cycle can be completed inside the nests. Lab experiments showed that females died sooner and invested less in reproduction in presence of red wood ants. Porcellio scaber rarely provoked an aggression response, but large numbers were carried as prey to the nest. These preyed isopods were mainly dried out corpses. Our results showed that the ant association incurred several costs for a facultative associate. Consequently, red wood ant nests and their surrounding territory act as an alternative habitat where demographic costs are offset by a stable resource provisioning and protection.

Abandoned Wood Ant Nests as Sites for Seedling Germination

We compared the species composition of seedlings germinated on ten recently abandoned Formica polyctena nests with the plant species on active nests and the plant species composition of the forest floor around the nests. Compared to no plants at all, when the nests were inhabited, seedlings of 12 different vascular plant species germinated on the nests after their abandonment. Four of them were myrmecochorous, of which two occurred only on the nests. After abandonment, ant activity ceases, the water content of the nests increases, and decomposition of the organic nest material is accelerated, resulting in faster nutrient mineralization. Consequently, more nutrients are available for plants. Our results suggest that abandoned nests can serve as “regeneration niches”, ensuring genetic diversity, especially in plant populations which rely mainly on vegetative propagation. However, more research on a larger scale is needed to assess the role of wood ants and their abandoned nests in forest vegetation development.

Assembly of a Hybrid Formica aquilonia × F. polyctena Ant Genome From a Haploid Male.

Formica red wood ants are a keystone species of boreal forest ecosystems and an emerging model system in the study of speciation and hybridization. Here, we performed a standard DNA extraction from a single, field-collected Formica aquilonia × Formica polyctena haploid male and assembled its genome using ~60× of PacBio long reads. After polishing and contaminant removal, the final assembly was 272 Mb (4687 contigs, N50 = 1.16 Mb). Our reference genome contains 98.5% of the core Hymenopteran BUSCOs and was pseudo-scaffolded using the assembly of a related species, F. selysi (28 scaffolds, N50 = 8.49 Mb). Around one-third of the genome consists of repeats, and 17 426 gene models were annotated using both protein and RNAseq data (97.4% BUSCO completeness). This resource is of comparable quality to the few other single individual insect genomes assembled to date and paves the way to genomic studies of admixture in natural populations and comparative genomic approaches in Formica wood ants.

Transfer of elements into boreal forest ants at a former uranium mining site

Ants can influence ecological processes, such as the transfer of elements or radionuclides, in several ways. For example, they redistribute materials while foraging and maintaining their nests and have an important role in terrestrial food webs. Quantitative data of the transfer of elements into ants is needed, e.g., for developing improved radioecological models. In this study, samples of red wood ants (genus Formica), nest material, litter and soil were collected from a former uranium mining site in Eastern Finland. Concentrations of 33 elements were analyzed by Inductively Coupled Plasma-Mass Spectroscopy/Optical Emission Spectroscopy. Estimated element concentrations in spruce needles were used as a proxy for studying the transfer of elements into ants via aphids because spruces host the most important aphid farms in boreal forests. Empirically determined organism/medium concentration ratios (CRs) are commonly used in radioecological models. Ant/soil CRs were calculated and the validity of the fundamental assumption behind the of use of CRs (linear transfer) was evaluated. Elements that accumulated in ants in comparison to other compartments were cadmium, potassium, phosphorus, sulfur, and zinc. Ant uranium concentrations were low in comparison to soil, litter, or nest material but slightly elevated in comparison to spruce needles. Ant element concentrations were quite constant regardless of the soil concentrations. Non-linear transfer models could therefore describe the soil-to-ant transfer better than conventional CRs.

Flow-on effects of an introduced tree species: Lodgepole pine plantation affects function and performance of boreal ants

Increased demand on forests to produce renewable biomass have accelerated the use of non-native tree species in forestry worldwide, often with negative effects on native biodiversity. In Sweden, the North American lodgepole pine (Pinus contorta Douglas ex. Loudon var. latifolia) was introduced already in the 1970’s. Even though the species now covers large areas, effects on native flora and fauna have not been thoroughly evaluated. In this study, we used a large-scale field experiment with 30 paired P. contorta and P. sylvestris stands in central and northern Sweden to study the effects of P. contorta plantation on the abundance and activity of red wood ants of the Formica rufa-group, density and species richness of other ant species, as well as possible effects on the abundance of the specialized ant predator Pella humeralis. We collected epigaeic insects with pit fall traps, surveyed wood ant mound density and volume, and monitored ant activity on trees in all stands. Our analyses revealed a trend with higher mound density in P. sylvestris compared with P. contorta stands, although wood ant worker catch did not differ between the pine species. However, red wood ant activity was significantly higher in P. sylvestris stands, suggesting that P. contorta is not a preferred tree species for the tending of aphids for honeydew. Total ant species richness as well as the catch of Camponotus herculeanus and Myrmica ruginodis were higher in P. sylvestris stands, suggesting that they constitute a more favorable habitat for ants. Ant species richness was negatively influenced by wood ants in P. sylvestris stands only. A similar pattern was shown for C. herculeanus. Myrmica ruginodis was more abundant in P. sylvestris stands only if no thinning had been applied. Wood ant catch was positively correlated with catch of the specialized predatory rove beetle Pella humeralis showing that there is a strong interaction between the predatory beetle and wood ants.Our results suggest that P. contorta plantations impact both red wood ants and associated species and although competition from wood ants is more pronounced in P. sylvestris stands they still constitute a more favorable habitat for most ant species. Thus, large scale plantation of the non-native but closely related P. contorta may negatively impact epigaeic assemblages, a pattern that is not compensated by released competition by dominant red wood ants.

Red wood ants shape epiphytic lichen assemblages in montane silver fir forests

The Formica rufa group comprises several ant species which are collectively referred to as “red wood ants” (hereafter RWA). These species have key roles in forest ecosystems, where they are ecologically dominant and greatly influence the dynamics of the habitat they colonise. Various studies have shown how their trophic activity may affect other organisms, which include both other invertebrates and plants. We can therefore hypothesize that their presence could affect the taxonomic and functional composition of epiphytes, despite clear information on such an effect is lacking. This study aimed to fill this research gap by evaluating whether the presence of red wood ants could affect the structure and composition of lichen communities. We selected two sites on the Apennine Mountains in Italy, where the red wood ant F. paralugubris was introduced from the Alps more than 50 years ago. In each site, lichen assemblages on Abies alba trees located within the colonised areas were compared to those from nearby, non-occupied areas. The results allowed for the identification of significant effects of F. paralugubris on the structure of lichen communities. Although there was no detectable impact on lichen species richness, a significant difference in their community composition between colonised and control sites was detected. Furthermore, ant presence seemed to be associated with specific lichen functional traits such as asexual reproduction. We argue that RWA could affect the lichen community either directly, e.g., by actively dispersing the species capable of asexual reproduction through their movements on trees (ant-mediated dispersion), or indirectly through herbivore exclusion. Finally, we also observed differences in β-diversity among the colonised and non-colonised sites.

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.

Biogeography and habitat preferences of red wood ants of the Formica rufa group (Hymenoptera: Formicidae) in Finland, based on citizen science data

Red wood ants (RWA) of the Formica rufa group are ecosystem engineers and important species in boreal and temperate forests. However, the permanent and temporal loss of forest habitats is a serious threat to their existence and is likely to increase with climate change. Due to the current threat of losing species, quick actions are needed. Reported here is the biogeography, relative abundance and habitat preferences of five species of RWA in Finland based on citizen science data. Species that occur in the lowlands of the Alps also occur throughout the southern parts of Finland. Only two of the five species, F. aquilonia Yarrow, 1955 and F. lugubris Zetterstedt, 1838, were common throughout the country, including northern Lapland. As their name suggests, RWAs occur mostly in forests and forest edges, but they also sometimes inhabit open or semi-open yards, mires and meadows. The most forest-specialist species is F. aquilonia and the least F. rufa Linnaeus, 1761. Surprisingly, the meadow wood ant F. pratensis Retzius, 1783 is the second most forest dwelling species, however, its preference for forest edges is clearly higher than for forest interiors. Based on current data, F. rufa may be the most tolerant of living close to buildings as its relative abundance in yards was much higher than that of the other species. The data obtained on distributions and relative abundances could be compared in the future with the results of similar surveys to detect changes in species distributions, relative abundances and habitat preferences.