Survival of transplanted nests of the red wood ant Formica aquilonia （Hymenoptera： Formicidae）： The effects of intraspecific competition and forest clear-cutting

The fitness and survival of ant colonies depend on the resources near their nests. These resources may be limited due to poor habitat quality or by intra- and interspecific competitions, which in extreme cases may cause the ant colony to perish. We tested the effect of intraspecific competition and habitat degradation （forest clear-cutting） on colony survival by transplanting 26 nests of the red wood ant （Formica aquilonia Yarrow, 1955） in 26 different forest areas that contained 0-11 conspecific alien nests per hectare. F. aquilonia is highly dependent on canopy-dwelling aphids, thus the removal of trees should cause food limitation. During the course of the 4-year experiment, 9 of the forests were partially clear- cut. We found that while forest clear-cutting significantly decreased the colonies＇ survival, intraspecific competition did not. As a highly polygynous and polydomous species, E aquilonia seems to tolerate the presence of alien conspecific colonies to a certain extent.     

Immune defense of ants is associated with changes in habitat characteristics

Although the immune functions of insects are known to correlate with body condition and food resources, the association between habitat structure and immune function is still largely unknown. We studied the effects of forest clear-cutting on encapsulation rate in gynes and workers in the forest-dwelling ant Formica aquilonia. Forest logging resulted in disturbed immunity in workers and gynes. Logging enhanced encapsulation reaction in gynes, whereas decreased that of workers. In gynes, there was a likely trade-off between growth and immune function that was apparent in terms of different investment in size and immune function in different habitats. In workers, however, such associations were not found. The results indicate that, because of disturbed immunity, environmental stress may increase susceptibility of wood ants to diseases and parasites in disturbed habitats.     

RETRACTED ARTICLE: Interspecific competition among ants in the boreal forest: Testing predictions from a linear hierarchical competition model

Competitive interactions often play an important role in local community structure, and particularly so in ant communities. We test predictions derived from a competition model proposed for Scandinavian ant communities by comparing ant nest densities inside and outside fifty randomly-selected territories of competitively dominant wood ants within 30 km of Ume?, northern Sweden. As predicted by the model, competitively-intermediate encounter species, as well as other territorial species, showed complementary occurrences with both studied wood ants, Formica aquilonia Yarrow and F. lugubris Zett.. In contrast, complementary abundances, i.e. lower density of competitively-inferior submissive species in the presence of wood ants, as predicted by the model, was not supported for all submissive species. Of the two studied wood ant species, the nest density of submissive species was negatively correlated only with F. aquilonia. Submissives as a group, as well as Myrmica-species, showed complementary abundances with F. aquilonia, but one Leptothorax and two Serviformica species had higher nest densities in the presence of this competitively-dominant species. We propose that, for Leptothorax, these deviations from the model predictions may be because of limited niche overlap with dominant wood ants, a small worker force and a timid behaviour which does not elicit aggression in wood ants. For the two Serviformica species, we propose a combination of protection against social parasites, inter-specific social control performed by F. aquilonia, and dominance relationships between competitively-inferior submissive species as reasons for the higher nest density inside F. aquilonia territories. Monogyny, and thereby smaller nests, lower worker force, smaller territory (as shown by this study) in F. lugubris, as compared to the polygyny in F. aquilonia, may also help explain the differences in their effects on subordinate species. Our analyses indicate that the linear competition hierarchy model proposed for Scandinavian ants accurately predicts the outcome of interaction between and community composition for dominant territorial and encounter species, but that it needs refinement with respect to the relationship between territorial and submissive species and the resulting community composition. Further studies are needed, especially addressing the complex relationships between these latter groups, and the effects of different competitively-dominant wood ants, to determine the mechanisms determining the outcome of these relationships and to more accurately predict community composition. Received 11 June 2007; revised 11 September 2007; accepted 17 September 2007.     

Habitat-related aggressive behaviour between neighbouring colonies of the polydomous wood ant Formica aquilonia

Ant species of the late succession stages usually have several queens and a polydomous colony structure, where several colonies occupy the same territory without competing. The wood ant Formica aquilonia is a good example of such a species in boreal forests. In this species, the lack of intraspecific competition may be caused by the stable environment and abundant food resources (e.g. excretions of tree-living aphids). We studied how habitat destruction, in the form of clear-cutting, affects aggressiveness between neighbouring colonies in F. aquilonia. Intercolonial relationships were more hostile in clear-cuts than in forest interiors. This aggression may be the result of increased intraspecific resource competition or alienation of neighbouring colonies caused by the loss of visual orientation cues. We suggest that the original polydomous relationships between colonies decline and colonies may start aggressive competition for the remaining resources.     

Wood ant colouration as an ecological indicator for the level of disturbance in managed coniferous forests

The search for ecological indicators of population well-being in natural and managed ecosystems is a crucial aspect of effective biomonitoring, conservation and nature protection. In long-term monitoring programs environmental stress has a measurable effect on naturally selected traits, such as body shape or size. However, changes in colouration provide information about early warning responses. The red wood ant Formica aquilonia is ecologically and territorially dominant among wood ant species in European boreal coniferous forest, and possesses variable red-brownish melanin-based cuticular colouration. F. aquilonia gynes, which are unfertilized queens, exhibit colour variability on the head, propodeum and abdomen, yet only head colour features allow setting clearly visible and symmetrical (left/right) classes of morphs. We studied phenotypic colour variability, melanisation and fluctuating asymmetry of colour patterns in faces of F. aquilonia gynes from natural (forest interiors) and disturbed (clear-cut zones and forest edges) habitats in Finland. We defined five variable, clearly visible and symmetrical (left/right) colour morphs of gynes' faces. Individuals of totally dark morph were present only in disturbed habitats. General analyses of melanisation degree showed, that gynes from disturbed habitats were significantly darker compared to those from forest interiors. Individuals from the same nest tend to have a similar degree of melanisation. The mean darkness of face was highest among individuals from forest-edge nests and the lowest from forest interior nests. In all habitat types the darkness of face increased with an increase in head width. Increase of cuticular melanisation in F. aquilonia gynes in disturbed habitats could be explained with the theory of thermal melanism and stress-induced immune defence. Although some amount of fluctuating asymmetry among left/right symmetrical colour variations on faces of F. aquilonia gynes was found, it was affected neither by habitat type nor by head width. Melanisation degree of red wood ants have a potency to be used as ecological indicator for the level of disturbance in managed coniferous forests and fluctuating asymmetry of colour variations in ants might be studied furtherly in cases of more severe environmental concerns.     

Response of the foraging behaviour of red wood ants (Formica rufa group) to exclusion from trees

Abstract 1 Because of the large numbers within a colony and their aggressive nature, red wood ants (Formica rufa group) have a potential to greatly influence the cold‐temperate forest ecosystem. Wood ants are omnivorous and hunt in trees as well as on the forest floor. 2 A field experiment in a mixed forest in central Sweden was carried out to examine (i) the foraging behaviour of wood ants on the forest floor and (ii) the impact of increased numbers of wood ants on the soil fauna. The foraging behaviour of wood ants was manipulated by excluding the ants from their food resources in the tree canopy, with the intention to increase ant activity on the forest floor. To estimate this activity, the number of trees with foraging ants, the numbers of ants going to and from their nests and the prey carried by home‐running wood ants were determined during the summer period. Pitfall traps were placed in the soil to determine effects on mobile soil invertebrates. 3 When excluded from local trees, wood ants searched other trees further away from the nests rather than searching more intensively for prey on the forest floor. By contrast to the initial hypothesis, more soil‐living prey were caught by ants in the control plots than in the plots where the local trees were not accessible to the wood ants. The proportion of soil‐living to tree‐living prey tended to be greater in the control plots. 4 In the treated plots (no access to the trees), wood ants had a negative effect on the activity of Linyphiidae spiders. There was little effect of wood ants on other soil invertebrates. 5 This study suggests that the role of wood ants as top predators in the forest soil food‐web in central Sweden is limited.     

Distribution of ant species and mounds (Formica) in different-aged managed spruce stands in eastern Finland

Mound‐building ants (Formica spp.), as key species, have large impacts on organisms and ecosystem functions in boreal Eurasian forests. The density, sizes and locations of ant mounds determine the magnitude and the spatial distribution of ant activities in forest ecosystems. Clear‐cutting can destroy wood ant colonies, and the species, abundance, dimensions and locations of ant mounds may change as forest stand structure changes with stand age. We compared ant species composition, ant mound numbers and dimensions, and the spatial distribution of mounds in Norway spruce [Picea abies (L.) Karst.] stands of different age (5, 30, 60 and 100 years) in eastern Finland. The mound density of Formica aquilonia Yarr. was greater in the two oldest stand age classes, while most mounds of Formica rufa L., Formica polyctena Först., Formica lugubris Zett., Formica exsecta Nyl. and Formica pressilabris Nyl. were found in the two youngest age classes. The mean volume, the volume per area and height/diameter ratio of F. aquilonia mounds increased with stand age. In the oldest stand age class, mounds were slightly smaller in well‐lit locations than in shade and near stand edges than further from the edges indicating that new mounds are established in well‐lit locations. Similarly, the longest slopes of the mounds faced south, indicating the importance of exposure to the sun. F. aquilonia mounds were concentrated near stand edges, and the spatial distribution of the mounds was aggregated in some stands. At the ecosystem level, the aggregation of ant mounds near stand edges may increase the edge productivity, as mounds concentrate resources to the edges and release nutrients after abandonment.     

An experimental study of behavioral mechanisms of relationships between red wood ants and ground beetles

Behavioral mechanisms regulating the competitive relations between the red wood ant Formica aquilonia and the ground beetles Carabus regalis, Pterostichus melanarius, P. magus, P. oblongopunctatus, Harpalus smaragdinus, and Amara nitida were studied by artificially initiated collisions between living insects as well as with the use of imitation models. Members of different functional groups within an ant family (aphid-milkers, hunters, and guards) behaved differently towards beetles. Active ants were shown to respond selectively to different features of the possible competitors, such as coloration, the presence of “appendages” (legs, antennae), body symmetry, rate of movement, and scent. Field and laboratory experiments demonstrated the ability of beetles to avoid collisions with active ants. The scent of anthill material attracted ground beetles, which consumed dead ants. The gained individual experience may allow the beetles to use supplementary forage resources in the territories controlled by ants. The flexible tactical patterns facilitate spatial segregation of ground beetles and ants in the same territory and result in a more complete utilization of food resources.     

Preadaptation of <Emphasis Type="Italic">Formica aquilonia</Emphasis> and <Emphasis Type="Italic">Formica lugubris</Emphasis> ants (Hymenoptera,Formicidae) to low wintering temperatures in Northeast Asia

The ants Formica aquilonia and F. lugubris which inhabit the entire forest zone of the North Palaearctic and are absent from the basins of the Yana, Indigirka, and Kolyma rivers were found in the coastal area of the Sea of Okhotsk. A possible climatic conditionality of their occurrence in the Northeast is considered based on the data on the biotopic distribution of ants, the temperature causation of their overwintering, and cold hardiness. On the Sea of Okhotsk coast, these ants overwinter at a depth of 40–200 cm in the soil. During winter, the minimum soil temperature at a depth of 40 cm under the anthill was ?5°C. The supercooling temperature of F. aquilonia was not lower than ?20.2 ± 0.5°C, that of F. lugubris, not lower than ?19.6 ± 0.4°C. Half of F. aquilonia individuals did not survive the daily exposure at ?13°C, F. lugubris, at ?16°C. These two cold-resistant species could inhabit some biotopes of the Kolyma River basin, similar to F. exsecta, F. lemani, and F. sanguinea, but they are absent there for some reasons that are not related to the temperature. A similar cold hardiness is characteristic of F. aquilonia in Estonia (Maavara, 1971, 1985), where it represents a side effect of diapause, since excessive cold hardiness has no adaptive significance for insects overwintering in the non-freezing soils of Estonia. Colonization of Siberia by ant species turned out to be possible only due to the existing cold hardiness, i.e. preadaptation to low temperature. On the Sea of Okhotsk coast, cold hardiness of the ants is non-adaptive due to the relatively mild conditions during winter.     

Genetic differentiation of disjunct populations of the antsFormica aquilonia andFormica lugubris in Europe

Summary The speciesFormica aquilonia andF. lugubris of the mound-building red wood ants have a disjunct boreoalpine distribution in Europe. The populations ofF. aquilonia in Finland, Switzerland and the British Isles show little genetic differentiation, whereas the populations ofF. lugubris show considerable differentiation. The Central European populations morphologically identified asF. lugubris can be genetically divided into two groups (here called types A and B). Type B is found in the Alps and the Jura mountains, and is genetically inseparable fromF. aquilonia. Type A lives sympatrically with type B in the Jura mountains and is also found in the British Isles. Sympatry of the two types in the Jura shows that these are separate species. It remains open whether type B is morphologically atypicalF. aquilonia or whether it is a separate species, perhaps with a past history of introgression betweenF. aquilonia andF. lugubris. The gene frequencies in the Finnish populations ofF. lugubris differ from those of both types A and B. Genetic differences withinF. lugubris indicate that the populations have evolved separately for a long time. The social structure ofF. lugubris colonies also shows geographic variation. The nests in Finland and the British Isles seem to be mainly monogynous and monodomous, whereas the nests in Central Europe are polygynous and form polydomous colonies.F. aquilonia has polygynous and polydomous colonies in all populations studied.     

The fate of an intentional introduction of Formica lugubris to North America from Europe

Red wood ants (Formica s.str.) are not prevalent in the forests of North America, but commonly occur in conifer and mixed conifer forests in northern Europe and Asia. In 1971, a European red wood ant species, Formica lugubris, was intentionally established in a 35‐year‐old predominantly mixed conifer plantation approximately 30 km north of QC, Canada. The purpose of its introduction was to evaluate the potential of this species as a biological control agent against conifer‐defoliating Lepidoptera species. This red wood ant introduction was monitored periodically for about 5 years after establishment, but its long‐term fate has not been reported. We visited this field site in 2005 and found that this species was well established, and we could locate some of the nests that resulted from the original release. We mapped and measured over 100 nests around the site of original release, which ranged from 5 cm in height to over 1 m. We estimated the population of introduced ants to have grown to over 8 million in the last 34 years. Significant clustering of nests suggests that these nests may be one supercolony. F. lugubris has become a dominant understory arthropod in this mixed forest, and is likely to have ecological impacts, including effects at the community and ecosystem level.     

Trophic position and seasonal changes in the diet of the red wood ant Formica aquilonia as indicated by stable isotope analysis

1. Red wood ants are among the most numerous generalist predators and strongly affect the composition of arthropod communities in forest ecosystems. However, their trophic position remains poorly understood. Stable isotope analysis was applied to study the trophic position of Formica aquilonia and reveal seasonal changes in its trophic links with both myrmecophilous aphids and other invertebrates in a mixed forest of western Siberia. 2. The δ¹⁵N values of F. aquilonia exceeded those of herbivores and aphids by approximately 3.5‰. Despite obligate trophobiotic relationships with aphids, F. aquilonia occupied the trophic position of first‐order predator. The higher content of ¹³C in the worker ants, compared with members of grazing food chains, was explained by their consumption of ¹³C‐enriched aphid honeydew. 3. Myrmecophilous tree‐dwelling aphids were enriched in ¹³C and ¹⁵N relative to grass‐inhabiting species, and the honeydew of tree‐dwelling aphids had higher δ¹³C values than those of the honeydew of grass‐inhabiting aphids. 4. The decrease in δ¹³C values of the worker ants from spring and summer to autumn apparently reflected the transition from the collection of tree sap and feeding on the aphid honeydew from trees with high ¹³C content in the spring and early summer to a more diverse liquid diet in late summer, which included ¹³C‐depleted honeydew of aphids from herbs. 5. The prevalence of the ¹⁵N‐depleted aphid honeydew in the ants' diet in the second half of the summer is discussed as one possible explanation for the seasonal decline in δ¹⁵N values of the worker ants.     

Respiration of wood ant nest material affected by material and forest stand characteristics

We studied differences in respiration of materials from different parts of wood ant nest (top, bottom, and rim) and from the nest surroundings (humus layer and mineral soil). Samples were taken from 8 wood ant (Formica aquilonia) nests in each of the two types of forest (birch and pine) in eastern Finland. The differences were related to material and forest stand characteristics (i.e., moisture, pH, carbon content, and C:N ratio). As a result, the highest respiration per g DW was measured at the top of ant nests in the birch forest. However, respiration did not significantly differ between the parts of ant nests in the pine forest. Respiration of the humus layers in both forest stands was on average higher, whereas respiration of the mineral soils in both forest stands was lower in comparison with respiration of the nest materials. The respiration per g C did not show any significant differences between different parts of nests and surrounding soil. The most important factors influencing respiration of the materials appeared to be moisture, carbon content, and pH. In conclusion, respiration of wood ant nest material is affected by the specific material and forest stand characteristics.     

An innate template for enemy recognition in red wood ants

The hypothesis of the innate template for perception and recognition of the enemy image in red wood ants Formica aquilonia Yarr. was tested by initiating conflicts between ants and predatory ground beetles, their competitors for space. Live beetles and their models with different characters were used. In nature, ants respond selectively to such features of competitors as dark coloration, the presence of “outgrowths” (legs, antennae), body symmetry, the rate of movement, and scent. Comparison of behavior of ants from natural colonies and “naive” (laboratory reared) ones showed that ants having no experience of encounters with competitors responded aggressively to an integral and sufficiently realistic enemy image. This suggests that red wood ants possess an innate template for recognition of potential competitors. At the same time, the ability to single out the key features and complete the integral image seems to require accumulation of experience.     

Effects of ants on the foraging of birds in spruce trees

I experimentally excluded ants from randomly selected spruce trees Picea abies near colonies of the wood ant Formica aquilonia. Foraging activity of birds in these trees was then compared to the foraging activity of birds in neighboring spruce trees, where ants were allowed to continue foraging. Birds which foraged in the foliage showed the effects of competition with ants: they visited the trees without ants more frequently, and for longer periods. In addition, the insects and spiders that they utilized as food were more abundant in the foliage of trees without ants. Cone-foraging birds, however, which fed on seeds in cones at the tops of the trees, did not show a preference for trees without ants. The differences of tree usage between foliage-gleaning and coneforaging birds can be explained by alteration of the birds' food supply by wood ants: ants did not feed on seeds in cones, and so did not compete with cone-foraging birds. However, foraging wood ants did feed on arthropods living in the foliage, thus reducing the amount of food available to birds there.     

Within territory abundance of red wood ants Formica rufa is associated with the body condition of nestlings in the Eurasian treecreeper Certhia familiaris

Studies on individual reproductive success in relation to interspecific competition between distantly related taxa are scarce. We studied whether the abundance of red wood ants Formica rufa -group is related to the breeding habitat selection, fecundity and offspring quality in the Eurasian treecreeper Certhia familiaris, an old-growth forest passerine. The nest-box occupancy data were gathered over a five-year study period, whereas the breeding performance analyses were based on a two-year data set. The abundance of wood ants, measured within 50  m around the nest-boxes, was not related to nest-box occupancy rate, fecundity or the physiological stress of nestlings. In contrast, the abundance of wood ants was negatively related to the offspring quality and life-history traits, such as lowered body mass, subcutaneous fat reserves, and tarsus length at fledging. Our results suggest that exploitative competition between distantly related taxa may have considerable and adverse influences on nestling quality as measured by body mass, subcutaneous, fat and tarsus length. Red wood ants may decrease the fitness of treecreepers as the lower body condition of nestlings has the potential to impair recruitment into the breeding population and, additionally, impair the future reproductive effort.     

Effects of overwintering temperature on the survival of the black garden ant (Lasius niger)

The overwintering temperatures of ants might well be elevated due to climate change. We studied whether the overwintering temperature affects the survival of the queens and whole colonies of the black garden ant, Lasius niger (Linnaeus, 1758). In two consecutive years (2009, 2010) we collected mated, colony founding queens (n=280) from the urban area of Turku, Finland. Half of the queens overwintered in +7 to +8 °C and the other half in +2 °C. After the overwintering period, we determined their survival rate and measured the body fat content, body size and immune defence (encapsulation rate) of overwintering queens. Using the same setup, we studied the survival of 1-year-old L. niger colonies (queen & workers). Overwintering at a lower temperature (+2 °C) decreased the survival of workers. The survival of colony founding queens differed between years, but unlike with workers, the overwintering temperature did not affect their survival: neither in the colony experiment nor in the single queen experiment. All of the surviving queens managed to produce their worker offspring at the same rate. The relative amount of body fat of queens was higher for those who overwintered at a lower temperature, which is likely a result of lower energy consumption. We did not detect differences in the encapsulation rate between the temperature treatment groups. The ability of colony founding queens to tolerate wide overwintering temperature variations present in urban environments may explain the success of the colony in urban areas. As the colony grows, the overwintering chambers may extend more deeply into the ground. Thus, workers may not have to cope with such cold conditions as colony founding queens.     

Nest mounds of red wood ants (Formicaaquilonia): hot spots for litter-dwelling earthworms

A previously undocumented association between earthworms and red wood ants (Formicaaquilonia Yarr.) was found during an investigation of the influence of wood ants on the distribution and abundance of soil animals in boreal forest soil. Ant nest mounds and the surrounding soil of the ant territories were sampled. The ant nest mound surface (the uppermost 5-cm layer) harboured a much more abundant earthworm community than the surrounding soil; the biomass of the earthworms was about 7 times higher in the nests than in the soil. Dendrodrilusrubidus dominated the earthworm community in the nests, while in soils Dendrobaenaoctaedra was more abundant. Favorable temperature, moisture and pH (Ca content), together with abundant food supply (microbes and decomposing litter) are likely to make a nest mound a preferred habitat for earthworms, provided that they are not preyed upon by the ants. We also conducted laboratory experiments to study antipredation mechanisms of earthworms against ants. The experiments showed that earthworms do not escape predation by avoiding contact with ants in their nests. The earthworm mucus repelled the ants, suggesting a chemical defence against predation. Earthworms probably prevent the nest mounds from becoming overgrown by moulds and fungi, indicating possible mutualistic relationships between the earthworms and the ants. Received: 21 November 1996 / Accepted: 3 April 1997     

Evidence for collective medication in ants

Social organisms are exposed to many pathogens, and have evolved various defence mechanisms to limit the cost of parasitism. Here we report the first evidence that ants use plant compounds as a collective mean of defence against microorganisms. The wood ants Formica paralugubris often incorporate large quantities of solidified conifer resin into their nests. By creating resin‐free and resin‐rich experimental nests, we demonstrate that this resin inhibits the growth of microorganisms in a context mimicking natural conditions. Such a collective medication probably confers major ecological advantages, and may be an unrecognized yet common feature of large, complex and successful societies.     

Widespread hybridization within mound‐building wood ants in Southern Finland results in cytonuclear mismatches and potential for sex‐specific hybrid breakdown

Hybridization and gene flow between diverging lineages are increasingly recognized as common evolutionary processes, and their consequences can vary from hybrid breakdown to adaptive introgression. We have previously found a population of wood ant hybrids between Formica aquilonia and F. polyctena that shows antagonistic effects of hybridization: females with introgressed alleles show hybrid vigour, whereas males with the same alleles show hybrid breakdown. Here, we investigate whether hybridization is a general phenomenon in this species pair and analyse 647 worker samples from 16 localities in Finland using microsatellite markers and a 1200‐bp mitochondrial sequence. Our results show that 27 sampled nests contained parental‐like gene pools (six putative F. polyctena and 21 putative F. aquilonia) and all remaining nests (69), from nine localities, contained hybrids of varying degrees. Patterns of genetic variation suggest these hybrids arise from several hybridization events or, instead, have backcrossed to the parental gene pools to varying extents. In contrast to expectations, the mitochondrial haplotypes of the parental species were not randomly distributed among the hybrids. Instead, nests that were closer to parental‐like F. aquilonia for nuclear markers preferentially had F. polyctena's mitochondria and vice versa. This systematic pattern suggests there may be underlying selection favouring cytonuclear mismatch and hybridization. We also found a new hybrid locality with strong genetic differences between the sexes similar to those predicted under antagonistic selection on male and female hybrids. Further studies are needed to determine the selective forces that act on male and female genomes in these newly discovered hybrids.