Ant assemblages in the taiga biome: testing the role of territorial wood ants

Summary Ants were collected with sets of pitfall traps in four coniferous-forest habitats in southern Finland. A three-level competition hierarchy concept was used to generate predictions on ant community structure. The levels of the hierarchy, and the respective predictions, from top to bottom were: (1) The dominant territorial wood ants (Formica rufa-group species), expected to exclude each other. (2) The other aggressive species, likely to be excluded by the F. rufa-group. (3) The submissive species, non-aggressive and defending only their nest, and thus likely to coexist with the dominants but in reduced numbers. As expected, the species of the F. rufa-group excluded each other, and the species number of the other aggressive ants was significantly cut down in the presence of the F. rufa-group. The aggressive species F. sanguinea and Camponotus herculeanus showed complementary occurrences with the F. rufa-group, and Lasius niger reduced occurrences. The number of the submissive species was not significantly affected by the F. rufa-group. However, pairwise correlation coefficients were significantly more often negative than positive between presence of the F. rufa-group and average proportion of pitfalls per set with a submissive species, each analyzed in turn. The result indicates that the F. rufa-group also reduced the colony densities of the submissive species. We conclude that in the taiga biome territorial wood ants are, after adjusting for physical vicissitudes of the environment, the major structuring force of ant species assemblages.     

Veränderungen in einem natürlichen vorkommen hügelbauender waldameisen derFormica rufa-Gruppe (Hymenoptera: Formicidae) im verlauf von drei jahren

Zusammenfassung In zwei vollständigen Aufnahmen während der Jahre 1966 und 1969 wurde das natürliche Vorkommen hügelbauender Waldameisen derFormica rufa-Gruppe in einem 1639,5 ha grossen Waldgebiet der Vorbergzone des Schwarzwaldes nahe Freiburg/Br. festgestellt. 1966 wurden 260 bewohnte Ameisenhügel gefunden; dies entspricht einer Ameisendichte von 15,9 Nestern je 100 ha. Bei der zweiten Aufnahme 1969 wurden in demselben Gebiet nur noch 208 bewohnte Nesthügel vorgefunden. Dieser im Verlauf von drei Jahren eingetretene Rückgang der Nestzahl um 24,4% hat die einzelnen Arten in unterschieldlichem Ausmass betroffen.Durch zusätzliche Registrierung aller während des Untersuchungszeitraums neu entstandenen, aber bereits wieder verlassenen Nesthügel bei der Zweitaufnahme konnte gezeigt werden, dass die im Verlauf von drei Jahren eingetretenen Veränderungen des Bestandes an bewohnten Ameisenhaufen weit übertroffen werden durch die Zahl nicht dauerhafter Neuanlagen.

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Summary A total census was carried out near Freiburg/Br. (Black Forest) in a woodland of 1639.5 ha (=4098.8 acres) in the year 1966 and again in 1969 to evaluate the number of natural populations of ants of theFormica rufa-group. In 1966, 260 lived-in ant hills were found, this corresponds to 15.9 nests per 100 ha (=6.3/100 acres). In 1969, only 208 lived-in ant hills were found in the same area. The single species were differently affected by this reduction of 24.4% in three years.The amount of change in the natural populations of red wood ants in the area studied is in fact very much higher than indicated by the reduction in numbers of lived-in ant hills. This could be evaluated by an additional registration of all «dead» nests during the second survey; a comprehensive list of the shifting in nesting sites and populations during these three years is given.

     

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.     

Abundance and distribution of organic mound-building ants of the Formica rufa group in Yellowstone National Park

Red wood ants (Formica rufa group) are ubiquitous in many conifer and mixed‐conifer forests of northern Europe and Asia. In contrast, relatively little is known about the abundance and distribution of the 24 North American F. rufa group species. As ants are important components of most soil invertebrate communities and are considered ecosystem engineers that alter the flow of energy and nutrients through terrestrial systems, it is important to gain information on their distribution and abundance. We conducted a survey for red wood ant mounds in Yellowstone National Park (YNP), Wyoming/Montana, USA, where human disturbance has been kept to a minimum for over 130 years. We found a total of 85 red wood ant mounds (0.11 to 0.17 mounds/ha) on 327 km of roads and 180 km of the hiking trails we surveyed. The occurrence of ant mounds was higher then expected by random distribution at elevations between 1600 and 2400 m, annual precipitation of 250 to 760 mm, middle and late successional lodgepole pine, late successional Douglas fir forest and non‐forested grassland/sagebrush prairie vegetation. Additionally, mounds were clustered in gently sloped not north‐exposed locations and in areas that had not recently burned. Most of the mounds detected were inhabited by Formica obscuripes Forel, which occupied 94% of the mounds sampled. Based on a multi‐criteria binary Geographic Information System model that we developed, we found that ant mounds were to be expected with a high probability in less then 1% of the YNP area. These results together with the detected low density and small size of the red wood ant mounds within the study area suggest that these insects have a much lower impact on invertebrate biodiversity and ecosystem processes, such as forest productivity and carbon and nutrient cycling on the ecosystem scale compared with their counterparts in European or Asian systems.     

Wood ants prefer conifers to broadleaved trees in mixed temperate forests

1. Wood ants, species of the subgenus Formica s. str., are one of the most important groups of insects in forest ecosystems due to their high abundance and activity.
2. We assessed the foraging pressure of Formica polyctena and Formica rufa, two dominant wood ant species in the Białowieża Forest. In addition, we compared coniferous and mixed stands in these respects.
3. In mixed stands F. polyctena visited more trees per plot than F. rufa. F. polyctena might forage a larger area due to higher population sizes, which further confirms higher foraging pressure of F. polyctena in comparison with F. rufa.
4. In our study area, host trees of the ant–aphid mutualism were larger in diameter than nonhost trees. The ants also preferred trees situated closer to their nests. In both mixed and coniferous stands, the most visited tree species was Norway spruce. Thus, we can conclude that Norway spruce seems to be preferred by the ant. The decline of living spruce in the Białowieża Forest may endanger their food source and impact the vitality of ant colonies.

     

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.     

Interference competition,not predation,explains the negative association between wood ants (Formica rufa) and abundance of ground beetles (Coleoptera: Carabidae)

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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     

Genetic differentiation in sympatric wood ants, <Emphasis Type="Italic">Formica rufa</Emphasis> and <Emphasis Type="Italic">F. polyctena</Emphasis>

Summary Direct observations have suggested that the closely related wood ants Formica polyctena and F. rufa represent different social organizations, with high queen number in F. polyctena and a high frequency of monogynous nests in F. rufa. We examined social organization and genetic population structure in a setup where populations of the two species are sympatric and gene flow between the species is possible. Our aim was to compare social organization in the species, and study evolutionary relationships between them. The observed relatedness among colony workers suggested that the difference in the level of polygyny is quantitative rather than qualitative, with a higher queen number in F. polyctena. The observed difference in polygyny was not accompanied by a difference in spatial genetic differentiation which was weak in both species. The genetic distance between the species is consistent with limited interspecific gene flow. Identification of a few possible F. rufa migrants in F. polyctena populations suggests potential interspecific gene flow. Thus, reproductive isolation of the species may not be complete when they are sympatric.Received 14 March 2003; revised 10 October 2003; accepted 20 October 2003.     

Population genetics of Formica aquilonia wood ants in Scotland: the effects of long-term forest fragmentation and recent reforestation

Formica aquilonia wood ants are forest specialists which play a key role in the ecology of forests in Europe. Many of the Scottish populations at the edge of the species distribution range occur in highly fragmented landscapes. We used ten microsatellite loci to study the genetic diversity and structure of populations from two contrasting regions (Inverpolly and the Trossachs) to set the Scottish populations in the context of conspecific populations in mainland Europe. Historically, both study regions have experienced extreme habitat loss and fragmentation over several centuries. Inverpolly has remained fragmented whereas large scale reforestation over the last century has greatly increased the forested area in the Trossachs. Despite the long history of fragmentation, genetic diversity in the Scottish populations was greater than in the populations in mainland Europe. Genetic diversity was similar in the two Scottish regions and no evidence of inbreeding was detected. However, the populations in Inverpolly showed more evidence of genetic bottlenecks, possibly due to more frequent stochastic events such as moorland fires. The ant populations in individual forests were genetically distinct and we detected no contemporary gene flow between forests. The most intensively studied forest where non-native conifer plantations now occupy the matrix between the remaining ancient woodland fragments showed evidence that admixture and gene flow between nests was reducing the past differentiation. This may reflect a dynamic response to the reconnection of previously isolated populations in forest fragments by recent reforestation.     

Long-term impact of agriculture on the survival of wood ants of the <Emphasis Type="Italic">Formica rufa</Emphasis> group (<Emphasis Type="Italic">Formicidae</Emphasis>)

The impact of agriculture on wood ants of the Formica rufa group was investigated in a small-scale agricultural landscape with many woodland fragments in the east of the Netherlands. An inventory of nests was carried out in 1986, and repeated in 2014. The number of nests of F. rufa and F. polyctena had drastically decreased during the intervening 28 years, especially in small patches of woodland bordering a maize field. The species F. rufa, which mainly disperses by flight of the queen, seems to be better adapted to habitat fragmentation than F. polyctena, where dispersal is mainly by colony splitting. However, the flying strategy can only be successful in a landscape where most woodland fragments have a good quality habitat; in the study area, the habitat quality had deteriorated. This is ascribed to the development of a tall, nitrophilous vegetation, a result of considerable nitrogen deposition from the atmosphere and the spraying of manure. The regional survival of wood ants in the east of the Netherlands is at risk in the agricultural landscapes we studied. This is due to the increasing probability of colonies becoming extinct and the decreased probability of a queen colonizing habitat patches. However, the negative influence of agriculture on the edge vegetation of patches of woodland can be decreased by leaving a wide strip between the arable land and the woodland that receives no manure.     

Chronicle of an impact foretold: the fate and effect of the introduced <Emphasis Type="Italic">Formica paralugubris</Emphasis> ant

Starting in 1958, red wood ants (Formica rufa group) from the Alps were transplanted to several Apennine forests along the Italian peninsula to be employed as biological control agents for tree insect pests. In the Campigna Biogenetic Nature Reserve, central Italy, hundreds of mounds of the dominant Formica paralugubris were repeatedly introduced, creating several populations that still survive today. In this study, we analyzed the temporal dynamics and the ecological impact of five of these populations. Their present state was assessed by censusing the total number of nest mounds and their volumes, while past changes were reconstructed from literature records. We also evaluated the impact of this species on autochthonous arthropod communities by comparing impacted and non-impacted areas and performing predation experiments. The density of nests and their volume remained stable or declined for a few years after transplant, and then they began to grow steadily. Local arthropods were severely affected, since almost all collected groups were significantly less represented in impacted than in non-impacted sites. Additionally, experiments with live bait demonstrated that potential prey have a significantly greater probability of being consumed in the areas occupied by F. paralugubris. These results prompt a thorough assessment of the fate of the introduced red wood ant populations, since their role as biological control agents has to be traded against the ecological impact on native arthropod communities. This is particularly relevant for highly biodiverse areas, such as the Campigna forest, that are home of several invertebrate species with conservation interest.     

Conservation genetics of the wood ant, Formica lugubris, in a fragmented landscape

Various intrinsic factors connected to the special features of sociality influence the persistence of social insect populations, including low effective population size, reduced amount of genetic variation easily leading to inbreeding depression, and spatially structured populations. In this work, we studied an isolated, small and fragmented population system of the red wood ant Formica lugubris, and evaluated the impact of social and genetic population structure on the persistence and conservation of the populations. The effective population size was large in our study population because all nests were polygynous. As a result, and despite the apparent isolation, the amount of nuclear genetic variability was similar to that in a nonisolated population system. Lack of inbreeding, as well as a high level of variability, indirectly suggests that this population does not suffer from inbreeding depression. The spatial distribution of genetic variation between local populations suggests intensive, but strongly male-biased, nuclear gene flow. Thus, the persistence of this population system does not seem to be threatened by any immediate social or genetic factor, but colonization of new habitat patches may be difficult because of restricted female dispersal.     

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.     

Pollen analysis and planted ancient woodland restoration strategies: a case study from the Wentwood, southeast Wales, UK

This paper considers the contribution of pollen analysis to conservation strategies aimed at restoring planted ancient woodland. Pollen and charcoal data are presented from organic deposits located adjacent to the Wentwood, a large planted ancient woodland in southeast Wales. Knowledge of the ecosystems preceding conifer planting can assist in restoring ancient woodlands by placing fragmented surviving ancient woodland habitats in a broader ecological, historical and cultural context. These habitats derive largely from secondary woodland that regenerated in the 3rd–5th centuries a.d. following large-scale clearance of Quercus-Corylus woodland during the Romano-British period. Woodland regeneration favoured Fraxinus and Betula. Wood pasture and common land dominated the Wentwood during the medieval period until the enclosures of the 17th century. Surviving ancient woodland habitats contain an important Fagus component that probably reflects an earlier phase of planting preceding conifer planting in the 1880s. It is recommended that restoration measures should not aim to recreate static landscapes or woodland that existed under natural conditions. Very few habitats within the Wentwood can be considered wholly natural because of the long history of human impact. In these circumstances, restoration should focus on restoring those elements of the cultural landscape that are of most benefit to a range of flora and fauna, whilst taking into account factors that present significant issues for future conservation management, such as the adverse effects from projected climate change.     

Does ant predation favour leaf beetle specialization on toxic host plants?

Generalist predators are frequently seen as evolutionary forces that narrow the host range in herbivorous insects. Predators may favour specialization of herbivores on host plants containing toxic chemicals (which can be used by herbivores for their own defence) if host plant‐derived defences provide better protection from enemies than do autogenously produced defences. We compared the effectiveness of these two defensive strategies in the larvae of six species of leaf beetle (Chrysomelidae) against wood ants (Formica rufa group) in field experiments. Ants were more strongly repelled by larvae with host plant‐derived, salicylaldehyde‐containing secretions than by larvae with various autogenous secretions, but collectively foraging ants ultimately overcame any type of chemical defence by social interactions, chemical signalling, and olfactory learning. As a result, ants killed all larvae of Chrysomela lapponica defended by salicylaldehyde‐containing secretions within 2 days of their introduction to willows within 15 m of ant nests. We conclude that in the field neither type of chemical defence provides complete protection against wood ants in the vicinity of their nests, and that evolutionary shifts from autogenous production of secretion to sequestration of plant allelochemicals in leaf beetles may be favoured mostly at low ant densities on the periphery of ant foraging areas.     

Independent hybrid populations of <Emphasis Type="Italic">Formica polyctena X</Emphasis><Emphasis Type="Italic">rufa</Emphasis> wood ants (Hymenoptera: Formicidae) abound under conditions of forest fragmentation

Combined genetic and morphological data indicate frequent hybridisation between the wood ants Formica polyctena Förster 1850 and F. rufa Linnaeus 1761 in Central Europe. The genetic and morphological traits give a concordant picture of hybridisation with a strong correlation between the genotypic admixture proportions at 19 microsatellite loci and the first vectors of a principal component analysis (P < 0.001) and of a 3-class discriminant analysis (P < 0.001) of 15 quantitative morphological characters. This integrative approach enabled a grouping into F. polyctena, the hybrid and rufa. Genetic differentiation between the hybrid and F. rufa is significantly larger than between the hybrid and polyctena, indicating gene flow mainly between the latter entities. A suggested gene flow bias towards F. polyctena agrees with differential queen acceptance and mating behaviour. Both genetic and phenotypic colony parameters indicate predominance of monogyny in F. rufa but of polygyny in polyctena and the hybrid. Hybrids are intermediate between the parental species in body size, diagnostic morphological characters, monogyny frequency, size of nest population, nest diameter and infestation rate with epizootic fungi. The three entities respond differently to woodland fragmentation. Hybrids are significantly more abundant in forests with a coherent area <300 ha than in woodland above this size. Regions with high hybrid frequency in Germany—the Eastern Oberlausitz (23%) and the Baltic Sea islands Darss, Hiddensee and Rügen (28%)—are characterised by a fragmented woodland structure whereas regions with low hybrid frequency—Brandenburg and the lower Erzgebirge (3.4%)—have clearly larger and more coherent forest systems. Data from other European countries indicate habitat fragmentation to be a facilitating factor but no essential precondition for interspecific hybridisation in these ants. Hybrids are hypothesised to have selective advantage in fragmented systems because of combining the main reproductive and dispersal strategies of the parental species.     

Genetic vulnerability of a remnant ant population

Ants are ecologically important species in many environments forming a big proportion of the total animal biomass. However, their special features such as sociality and complementary sex-determination system have received little attention in conservation discussions. We examined the social and genetic structure of the hairy wood ant Formica lugubris in Ireland to evaluate factors relevant to the vulnerability of small, isolated wood ant populations. We also clarified the conservation status of the species in Ireland. Our results showed that the populations are mainly monogynous (one queen per nest). Consequently the effective population size is very low (<100 individuals) in Ireland. We found extremely little genetic variation, signs of inbreeding and inbreeding depression, which can be a consequence of the small effective population size and of the restricted gene flow due to strong isolation of populations. Putatively high genetic diversity at the sex-determining locus can reflect a larger population in the past. The study shows that even though the population may seem stable because of the long life span of queen ants, sociality can impact species conservation by keeping the effective population size small. According to our results, the hairy wood ant can be considered native to Ireland. Hence it needs urgent protection and the genetic issues need to be considered in the future management strategies.     

Stand type is more important than red wood ant abundance for the structure of ground‐dwelling arthropod assemblages in managed boreal forests

• 1 The relationships between red wood ants (Formica rufa group) and other ground‐dwelling arthropods were studied in young managed forests stands in Eastern Finland. The main objectives were: (i) to test the influence of stand type (dominant tree species; age: sapling versus pole stage) and numbers of red wood ants on the occurrence of other ground‐dwelling arthropods and (ii) to study the occurrence of red wood ants versus other arthropods on a distance gradient from ant mounds. We used pitfall traps set in 5–14‐year‐old sapling stands and 30–45‐year‐old pole‐stage stands of Scots pine (Pinus sylvestris L.) and birch (Betula spp.) forests.
• 2 Pitfall trap catches of red wood ants did not vary significantly between the forest stand types, although some groups of other arthropods showed clear responses to stand type (e.g. catches of other Formicinae and Gnaphosidae were higher in sapling stands than in pole‐stage stands). The number of red wood ants clearly explained less of the variation in assemblages of other ground‐dwelling arthropods than the forest stand type.
• 3 Red wood ant numbers decreased significantly with distance from the mounds, but the other ground‐dwelling arthropods were insensitive to this gradient or even showed a preference for proximity to ant mounds and high ant activity.
• 4 The results obtained in the present study suggest that wood ants do not have strong effects on several other ground‐dwelling arthropod groups in young managed forests other than in the immediate vicinity of their mounds.

     

Immigration and formation of mixed colonies in red wood ants (Hymenoptera,Formicidae)

Long-term field studies of the composition and spatial structure of settlements of ants of the Formica rufa group were carried out in two regions of Russia (Moscow and Arkhangelsk provinces). Fragmentation of damaged nests followed by reintegration of the fragments is the main way of formation of mixed colonies of ants from different nests (including different species). The principal factor of nest fragmentation is their damage by wild boars, bears, and in some localities, by poachers. The formation of mixed nests and nest complexes with participation of different Formica species was observed. They are formed by joining the ants from several damaged nests or by a colony from a destroyed nest immigrating into an intact one. Regular damage of many nests leads to the formation of broad zones of mixed colonies. The mixed colonies including 2–3 species of wood ants have recently become common. The phenomenon of mixed colonies raises a question as to the relative importance of two basic principles (sociality and specific identity) in the life of ant societies and demonstrates the priority of the social principle.