Testate amoebae (Testacea) in a <Emphasis Type="Italic">Formica lugubris</Emphasis> nest: Fauna composition and structure

The fauna of testate amoebae (Testacea) in a Formica lugubris anthill and in litter layers from the control soil area in a spruce forest, Moscow region, was represented by a total of 43 species and subspecies. Their numbers ranged from 11000 ind./g air-dry substrate in the surface layer of the anthill to 62000 ind./g in the lower litter layer A0H/A1, with variation in species diversity between the samples being insignificant (24–30 species). Representatives of the genera Centropyxis, Cyclopyxis, Plagiopyxis, Corythion, and Trinema were dominant, whereas most of other species were few in numbers. The group of testacean species in the anthill was not specific, being a derivative of the testacean complex inhabiting the surrounding soil. Differences between litter layers and between these layers and substrates from the anthill concerned mainly the composition of the testacean community, which proved to change in the course of plant material decomposition: the aerophilic complex of upper layers (dominated by Centropyxis aerophila and Trinema lineare) was substituted by an edaphophilic complex (with Plagiopyxis declivis being dominant) in the nest mound of the anthill and in the lower litter layers. Plagiopyxis penardi was a eudominant species in the nest mound. Its especially high abundance is explained by deep transformation of plant remains in the nest (from moder to mull-like substrate) as the result of ant life activities.     

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.     

Population studies on hill building wood-ants of the Formica rufa-group

Summary Population studies on hill building wood-ants of the Formica rufa-group.Total inventories of inhabited ant hills of the Formica rufa-group were carried out in a part of the Black Forest amounting to 1,640 ha (4,100 acres) near Freiburg im Breisgau on five occasions during the period 1966–1978. Nest densities of 12.7 to 19.1 per 100 ha were recorded. Mean annual nest mortality for the three most abundant species (F. rufa, F. polyctena, F. pratensis) was 21–33% (Table 2); it remained almost the same during all intervals between inventories for all age classes studied (Fig. 2). The mean annual nest natality varied between 31 and 51%. It was therefore concluded that changes in the numbers of ant hills (Fig. 1) resulted from nest natality rather than from the reasonably stable nest mortality. Nest size was assessed by height and diameter. The development of nest size showed a marked increase with age for F. rufa and F. polyctena reaching a maximum at 2–5 years. Mean life expectancy of small nests was markedly lower than that of larger nests (Fig. 3). Consequently in the large size classes the numbers of old nests were significantly greater than the numbers of newly founded nests (Fig. 4). Results are compared with those recorded in the literature for palearctic and nearctic species belonging to the Formica rufa-group.Supported by Ministerium für Ernährung, Landwirtschaft und Umwelt Baden-Württemberg (MELU) and Gesellschaft zur Förderung der Forst- und Holzwirtschaftlichen Forschung Freiburg     

Polygyny reduces rather than increases nestmate discrimination cue diversity in Formica exsecta ants

Although the majority of social insect colonies are headed by a single queen, some species possess nests that contain numerous reproductive queens (polygyny), a trait that is particularly widespread amongst the ants. Polygyny is often associated with a lack of conspecific inter-nest aggression between workers. This is hypothesised to result from increased nestmate cue diversity within nests, since polygynous nests are more genetically diverse than monogynous nests. Alternatively, it may reflect the common origin of polygynous nests that form polydomous networks. We exploit the recent discovery that the nestmate discrimination system in the ant Formica exsecta is based on cuticular hydrocarbons to investigate cue (Z9-alkenes) diversity in several monogynous and polygynous populations. Contrary to previous predictions, in all polygynous populations, the variation between nests in the Z9-alkene profiles was reduced relative to that found in monogynous populations. However, nest-specific Z9-alkene profiles with little variation amongst nestmate workers were still maintained irrespective of nest type or population. This suggests a very effective gestalt mechanism that homogenises the chemical discrimination cues, despite genetic diversity within colonies. Although the reduction in variation between nests was associated with reduced worker aggression on the population level, it cannot totally explain the weak aggression associated with polygynous populations.     

Nesting preferences of the threatened wood ant <Emphasis Type="Italic">Formica exsecta</Emphasis> (Hymenoptera: Formicidae); implications for conservation in Scotland

Wood ants are a dominant and ecologically important component of northern coniferous forests with interactions at many trophic levels. Each species exhibits specific habitat preferences which need to be understood if conservation measures are to be successful. In Britain, the rare narrow-headed ant Formica exsecta has disappeared from much of its former range and is now largely restricted to the highlands of Scotland where it is found in open canopy woodland and along forest edges. Nest locale of one small and vulnerable population at the edge of its current range, were compared with those of random locations within the same habitat and with actual nests within the stronghold of the Abernethy-Glenmore complex in Strathspey. After data exploration with Redundancy Analysis, stepwise multiple regression was used to create a model which best estimated the variance in nest location using a parsimonious selection of vegetation and environmental variables. The input variables included were light, soil moisture, altitude, tree stature and distribution, vegetation structure and composition, and ground characteristics. F. exsecta clearly exhibited preferences for the position of nest mounds in relation to light, vegetation and tree cover. Forest location was also important in determining which variables nests were affected by. This study highlights the importance of maintaining a dynamic mosaic of different-aged woodland enabling early successional species such as F. exsecta, with suitable areas to move to as conditions change and allowing the co-existence of all wood ant species.     

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.     

Nest-building in ants <Emphasis Type="Italic">Formica exsecta</Emphasis> (Hymenoptera,Formicidae)

Representatives of the subgenus Coptoformica build composite aboveground nests in much the same manner as red wood ants do. However, despite the superficial similarity, Coptoformica nests show a number of substantial differences that impact on the way in which ants use the nest. The aboveground part of a Formica (Coptoformica) exsecta nest consists of the temporary layer, the outer crust layer, the mound, and the intermediate layer. The material of all the layers includes soil and fragments of plants (picked up or excised), the relative proportion of the two components differing among the layers. The thick, soil-rich crust layer acts as armature and allows a variety of anthill shapes. The intermediate layer and the upper part of the mound harbor quite a few large chambers. Building material is often transferred from nest to nest. On the whole, F. exsecta nests are plastic and movable, which facilitates the activity of these ants in ecotone biotopes.     

Yeast communities of <Emphasis Type="Italic">Formica aquilonia</Emphasis> colonies

Yeast abundance and species diversity in the colonies of Formica aquilonia ants in birch–pine grass forest near Novosibirsk, Russia, were studied. The average yeast number in the anthill material was 10³–10⁴ CFU/g, reaching 10⁵ CFU/g in the hatching chambers. Typical litter species (Trichosporon moniliiforme and Cystofilobasidium capitatum) were predominant in soil and litter around the anthills. Apart from these species, ascomycete species of the family Debaryomycetaceae, Debaryomyces hansenii, and Schwanniomyces vanrijiae were predominant in the anthill material. Yeast population of the ant’ bodies consisted exclusively of the members of the last two species. Thus, highly specific yeast communities formed in the colonies of Formica aquilonia ants differ from the communities of surrounding soil. These differences are caused by environment-forming activity of the ants.     

Formation of springtail (Collembola) communities during colonization of ant-hills

Formation of springtail assemblages in resettled nests of the ant Formica aquilonia was studied. The species composition and relative abundance of springtails was shown to depend on the age, hydrothermal regime, and condition of the ant nest. The abundance and species diversity of springtails increases during the initial period of the ant-hill growth. Old nests are characterized by the minimum species diversity, with one or two distinct dominants. High ecological plasticity, including resistance to high temperatures and low humidity, allows Lepidocyrtus violaceus to maintain its stable dominant position in the ant-hills. Species with relatively narrow hygro-and thermopreferenda quickly occupy temporal favorable niches, reach high abundance, and quickly disappear as the microclimatic conditions change.     

Comparative study of actinomycete communities associated with Lasius niger and Formica cunicularia ants and their nests

A comparative analysis of the abundance and biodiversity of actinomycete communities isolated from living ants Lasius niger and Formica cunicularia, as well as their anthills, has been carried out for the first time. The number of actinomycetes detected in L. niger ants is close to the number of actinomycetes in their anthills and one order is higher than that for F. cunicularia. Actinomycete communities of anthills and the intact soil are characterized by a high diversity, equitability, absence of severe dominants, and the presence of common species and differ by number and the range of species. Biodiversity of actinomycetes associated with living ants is considerably lower than in their nests and the surrounding soil and the range of actinomycetes is specific for both the species of ants.     

Effects of Formica podzolica ant colonies on soil moisture,nitrogen, and plant communities near nests

1. Ants are widely regarded as ‘ecosystem engineers’ because their nest construction and contributions to nutrient cycling change the biological, chemical, and physical properties of the soil around their nests. Despite increasing attention to ant manipulation of soil ecosystems, the extent to which many common species influence soil properties, as well as nutrient uptake and community composition of plants near nests, is still unknown. 2. This study tested hypotheses that activities of a common subalpine ant, Formica podzolica, alter soil moisture and pH, redistribute nitrogen around nests, and affect plant species abundance and ground cover. 3. A combination of field sampling techniques showed that distance from a nest had a positive relationship with soil moisture and a negative relationship with plant abundance next to and downhill from nests. Slope aspect also affected plant communities, with downhill transects having higher plant cover and above‐ground biomass than uphill transects. A stable isotope analysis did not reveal that plants near nests had enriched ¹⁵N, but there were substantial differences in ¹⁵N among sites. 4. Overall, this study uncovers significant impacts of F. podzolica on the subalpine microhabitats directly surrounding their nests.     

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.     

Polymorphic microsatellite DNA markers in the ant Formica exsecta

Highly polymorphic genetic markers provide a useful tool for estimating important genetic parameters in studies of the evolution of sociality in insects. Here we report 14 polymorphic microsatellite markers developed in the ant Formica exsecta. The number of alleles found ranged between 3 and 18 per locus. These markers were developed for studying genetic population structure and mating structure in F. exsecta populations with varying social organizations (monogyne and polygyne types of societies). Cross‐species amplification indicated that some of the markers might be usable even in species belonging to different subfamilies.     

Effect of host species,host nest density and nest size on the occurrence of the shining guest ant <Emphasis Type="Italic">Formicoxenus nitidulus</Emphasis> (Hymenoptera: Formicidae)

Understanding habitat requirements of species is important in conservation. As an obligate ant nest associate, the survival of the globally vulnerable shining guest ant, Formicoxenus nitidulus, is strictly tied to that of its hosts (mound building Formica ants). We investigated how host species, nest density, inter-nest distance and nest mound size relate to the occurrence of F. nitidulus. In total, 166 red wood ant nests were surveyed in SW Finland (120 Formica polyctena, 25 F. rufa, 14 F. aquilonia, 5 F. pratensis, and 2 F. lugubris). Overall, F. nitidulus was found in 60% of the nests. For the actual analysis, only F. polyctena and F. rufa nests were included due to the small number of other nests. F. nitidulus was more likely to be found among F. polyctena than F. rufa. Also, while inter-nest distance was not important, a high nest density, commonly found in polydomous (multi-nest) wood ant colonies, was beneficial for F. nitidulus. The guest ant was also more likely to be found in large host nests than small nests. Thus, our results show that the best habitat for the guest ant is a dense population of host nest mounds with a high proportion of large mounds. Conservation efforts should be directed at keeping the quality of the red wood ant habitats high to preserve their current populations and to increase colonization. This will not only benefit the guest ant, but also a plethora of other species, and help in maintaining the biodiversity of forests.     

Testacean communities in perturbed soils: the influence of the wandering albatross

On Île de la Possession (Crozet Archipelago, sub-Antarctica), the testate amoebae (Protozoa, Rhizopoda) fauna in soils around abandoned and occupied nests of the wandering albatross (Diomedea exulans) was investigated. A comparison with control samples, a cluster analysis and several ordination techniques indicated that the presence of the breeding albatrosses induced modifications in physico-chemical soil characteristics and in the soil inhabiting testacean fauna. Only 11 testate species occurred frequently in the soils in the albatross’ zone. Soils around occupied nests had significantly higher moisture values, less acid pH values, an increased specific conductance and elevated phosphate and ammonium concentrations. Highly influenced testacean communities were characterised by high abundances of Difflugiella oviformis and extremely high abundances of Trinema lineare, resulting in a very low diversity and evenness within these communities. The intermediary situation of one abandoned nest indicated that soils around abandoned nests evolve gradually back to undisturbed soils. Although an overall negative influence on the testacean diversity, a limited albatross’ influence may increase the living fraction within the testacean soil communities.     

Molecular markers allow sibling species identification in red wood ants (Formica rufa group)

Red wood ants (Formica rufa group) constitute a group of species that are considered to be among the most promising bioindicators in forest ecosystems. However, because of their morphological similarity and intraspecific variability, morphological species identification can be difficult. Considerable expertise is necessary to discriminate between the sibling species F. lugubris and F. paralugubris, two species that often live in sympatry in the same Alpine forests. New taxonomic tools providing rapid and reliable species identification are needed. We present a simple and reliable molecular technique based on mtDNA (COI gene) and a restriction enzyme for discriminating between F. lugubris and F. paralugubris. We confirm the validity of this method with a Bayesian analysis based on microsatellites. This new molecular tool represents a clear breakthrough for discriminating between F. lugubris and F. paralugubris and is likely to be helpful in large‐scale biomonitoring.     

Reproductive parameters vary with social and ecological factors in the polygynous ant Formica exsecta

Due to their haplo‐diploid sex determination system and the resulting conflict over optimal sex allocation between queens and workers, social Hymenoptera have become important model species to study variation in sex allocation. While many studies indeed reported sex allocation to be affected by social factors such as colony kin structure or queen number, others, however, found that sex allocation was impacted by ecological factors such as food availability. In this paper, we present one of the rare studies that simultaneously investigated the effects of social and ecological factors on social insect nest reproductive parameters (sex and reproductive allocation, nest productivity) across several years. We found that the sex ratio was extremely male biased in a polygynous (multiple queens per nest) population of the ant Formica exsecta. Nest‐level sex allocation followed the pattern predicted by the queen‐replenishment hypothesis, which holds that gynes (new queens) should only be produced and recruited in nests with low queen number (i.e. reduced local resource competition) to ensure nest survival. Accordingly, queen number (social factor) was the main determinant on whether a nest produced gynes or males. However, ecological factors had a large impact on nest productivity and therefore on a nest's resource pool, which determines the degree of local resource competition among co‐breeding queens and at what threshold in queen number nests should switch from male to gyne production. Additionally, our genetic data revealed that gynes are recruited back to their parental nests after mating. However, our genetic data are also consistent with some adult queens dispersing on foot from nests where they were produced to nests that never produced queens. As worker production is reduced in gyne‐producing nests, queen migration might be offset by workers moving in the other direction, leading to a nest network characterized by reproductive division of labour. Altogether our study shows that both, social and ecological factors can influence long‐term nest reproductive strategies in insect societies.     

Individual variation in immune function in the ant Formica exsecta; effects of the nest, body size and sex

It has been generally thought that sex differences in the immune system are the result of the differing life history strategies of the sexes, although the available data are not entirely consistent with the hypothesis. In this study, we studied the variation in the immune function in the mound-building wood ant Formica exsecta. F. exsecta has two forms of males, distinguished by size: the small males (micraners) and the large males (macraners), which die after the mating period, whereas females live tens of years laying their eggs. We found that in general males have a lower encapsulation response against nylon monofilament (i.e. lower immune function) than queens. Among males, the micraners had a lower encapsulation rate than the macraners. However, in queens, there was no correlation between size and encapsulation rate. The origin nest had an effect on the encapsulation rate of males: males from the large nests had a stronger encapsulation rate than males from small nests. However, in queens, nest size did not have any effect on encapsulation response. The observed variation between sexes and individuals in the encapsulation rate is discussed in the context of reproductive strategies and parasite-mediated sexual selection.     

The dynamics of vegetation on and around <Emphasis Type="Italic">Formica rufa</Emphasis> nests

The species composition and structure of plant communities related to the activity of Formica rufa ants were studied in green moss pine forests located in Nerusso-Desnyanskoe Polesye (Bryansk Province). Four types of microsites were established and examined: (1) the mound of an active ant nest, (2) the soil bank around an active ant nest, (3) the mound of an abandoned nest, and (4) the soil bank around an abandoned nest. The background plant community was considered as the fifth type. According to MRPP method, all pairs of microsites except (3–4) were different both in plant species composition and species abundance. The microsites created by Formica rufa increase the habitat capacity and β diversity of the plant communities due to the appearance of meadow species requiring richer soils.     

Colony success of the submissive ant Formica fusca within territories of the dominant Formica polyctena

Abstract. 1. Twenty-three nests of the submissive ant Formica fusca L. were sampled in two adjacent territories of the dominant wood ant Formica polyctena Först. The nests were dug up at different distances from the wood-ant mounds. Distance is assumed to be inversely related to the extent of disturbance of F. fusca by F. polyctena.
2. Colony success of F. fusca was assayed by counting the numbers of workers (colony size), and worker and sexual offspring in a nest. For individual size, the head width and dry weight of fifty workers per nest were measured.
3. Colony size correlated positively with the numbers of worker and sexual offspring in one territory, but only with worker offspring in the other.
4. Distance from the wood-ant mound correlated positively with colony size and numbers of worker and female offspring in one territory. In the other territory distance correlated positively with number of females, but negatively with number of worker offspring. No significant correlations between distance and the size measurements were obtained.
5. Because of its nonaggressive behaviour F. fusca may nest fairly close to a wood-ant mound but is likely to show reduced abundances in terms of nest density and forager number, and, ultimately, lowered fitness.