Impact of Formica exsecta Nyl. on seed bank and vegetation patterns in a subalpine grassland ecosystem

The mound building ant Formica exsecta Nyl. is widely distributed in grassland ecosystems of the Central European Alps. We studied the impact of these ants on seed bank and vegetation patterns in a 11 ha subalpine grassland, where we counted over 700 active ant mounds. The mounds showed a distinct spatial distribution with most of them being located in tall‐grass, which was rarely visited by ungulates (red deer; Cervus elaphus L.). Heavily grazed short‐grass, in contrast, seemed to be completely avoided by ants as only few mounds were found in this vegetation type. The species composition of the ant mound and grassland seed banks was quite similar, i.e. from 15 common plant species 12 were found in both seed bank types. We found the same proportions of myrmecochorous seeds in ant mound and grassland soil samples. In contrast, the number of seeds was 15 times higher in mound compared with the grassland soil samples. Also, the vegetation growing on ant mounds significantly differed from the vegetation outside the mounds: graminoids dominated on ant mounds, herbaceous and myrmecochorous species in the grassland vegetation. We found significant continuous changes in vegetation composition on gradients from the ant mound centre to 1 m away from the mound edge. Overall, F. exsecta was found to have a considerable impact on seed bank and vegetation patterns in the grassland ecosystem studied. These insects not only altered grassland characteristics in the close surrounding of their mounds, but also seem to affect the entire ecosystem including, for example, the spatial use of the grassland by red deer.     

Interactive effects of soil-dwelling ants, ant mounds and simulated grazing on local plant community composition

Interactions between aboveground vertebrate herbivores and subterranean yellow meadow ants (Lasius flavus) can drive plant community patterns in grassland ecosystems. Here, we study the relative importance of the presence of ants (L. flavus) and ant mounds under different simulated grazing regimes for biomass production and species composition in plant communities. We set up a greenhouse experiment using intact soil cores with their associated vegetation.We found that plant biomass production in the short term was affected by an interaction between simulated grazing (clipping) and ant mound presence. Clipping homogenized production on and off mounds, while in unclipped situations production was higher off than on mounds. During the experiment, these differences in unclipped situations disappeared, because production on unclipped mounds increased. Plant species richness was on average higher in clipped treatments and patterns did not change significantly over the experimental period. Plant community composition was mainly affected by clipping, which increased the cover of grazing-tolerant plant species. The actual presence of yellow meadow ants did not affect plant community composition and production.We conclude that the interaction between ant mounds and clipping determined plant community composition and biomass production, while the actual presence of ants themselves was not important. Moreover, clipping can overrule effects of ant mounds on biomass production. Only shortly after the cessation of clipping biomass production was affected by ant mound presence, suggesting that only under low intensity clipping ant mounds may become important determining plant production. Therefore, under low intensity grazing ant mounds may drive the formation of small-scale plant patches.     

Vegetation of ant-hills in a mountain grassland:effects of mound history and of dominant ant species

Vegetation in grasslands with well-developed long-lastingant-hills in the Slovenské Rudohorie Mts., Slovakia, was studiedin relation to (i) position on the mound, (ii) ant speciesforming the mound, and (iii) history of the mound. Permanent plotrecordings of mound size and dominant ant species started fifteen years priorthe study began provided information on the history of individual mounds.The mound vegetation bears a striking similarity to vegetation insimilar habitats across a large part of Europe due to presence of species suchas Agrostis capillaris, Dianthusdeltoides, Polytrichum commune agg.,Thymus pulegioides, and Veronicaofficinalis. Out of the three major ant species-groups presentat the site (Lasius flavus, Tetramoriumcaespitum and Formica spp.), L.flavus had the most pronounced and the most lasting effect on themound vegetation. The dominance of the plant species listed above increased withthe time span over which the mound was inhabited by L.flavus. The effects of other species on vegetation composition,though discernible from short-term observation, disappeared over severalyears. The mounds proper did not differ from the undisturbed grassland in theproportion of myrmecochorous plants or plants with specific seed size ordormancy type. However, there was a highly significant concentration ofmyrmecochorous plants in the grassland patches immediately neighbouring themounds; this is likely to be due to seeds deposited there by the workers fromthe nest after the elaiosomes had been consumed. The mound vegetation wascomposed mainly of species with long stolons or rhizomes; however, there was nosignificant difference in formation type or length of stolons/rhizomesbetween mounds and the rest of the grassland or among mounds formed by differentant species.     

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.     

Social structure in the ant Lasius flavus: multi-queen nests or multi-nest mounds?

Abstract.

• 1 The yellow subterranean ant Lasius flavus is generally considered to have a single queen in mature nests. Yet, many anecdotal observations have suggested that the conspicuous earth mounds of this ant may, at least occasionally, contain several queens and their offspring.
• 2 To estimate the number of matrilines per mound, samples for horizontal starch gel electrophoresis were collected from a total of thirty-seven mounds in three sites of old chalk-grassland in southern England.
• 3 Allozymatic variation at an esterase and hexokinase locus showed that probably more than 50% of the mounds contained more than one queen and that the mean number of queens per mound is at least 1.5–2.
• 4 Further results strongly suggested that workers from different matrilines may share the same galleries, at least in the top of the mound.
• 5 The apparently variable social structure of Lasius flavus colonies is discussed, both technically and with reference to ecological processes related to the presence of ants in the nest mounds.

     

Organic mound-building ants: their impact on soil properties in temperate and boreal forests

Ants are important components of most soil invertebrate communities, and can affect the flow of energy, nutrients and water through many terrestrial ecosystems. The vast majority of ant species build nests in the mineral soil, but a small group of ants in temperate and boreal forests of Eurasia and North America build large parts of their nests above‐ground using organic materials collected from the surrounding soil. Many studies have shown that ants nesting in mineral soil can affect water infiltration rates, soil organic matter (OM) content, and nutrient cycling, but much less is known on how mound‐building ants influence soil physical and chemical properties. In this paper we summarize what is known on the soil impacts of organic mound‐building ants in temperate and boreal forests, and how these ants could be affected by ecosystem disturbance and future climate change. Much of this information comes from studies on Formica rufa group ants in Europe, which showed that CO₂ emissions and concentrations of C, N, and P are usually higher in ant mounds than in the surrounding forest soil. However, ant mounds are a minor component of total soil C and nutrient pools, but they do increase spatial heterogeneity of soil water and available nutrients. Mound‐building ants can also impact tree growth, which could change the quantity and quality of OM added to soil. Forest management, fire, and projected climate change, especially in boreal forests, could affect mound‐building ant population dynamics, and indirectly, soil properties.     

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.     

Effect of three mound-building ant species on the formation of soil seed bank in mountain grassland

The effect of three ant species (Lasius flavus, Formica spp., Tetramorium caespitum) on soil seed bank formation was studied in temperate mountain grassland. Seed removal experiments, analysis of soil seed content and seed survival experiments were carried out to evaluate the influence of ground ants on the seed fate. In the seed removal experiment seeds of 16 species, including 5 species with elaiosome-bearing seeds (myrmecochores), were exposed and their removal followed for 39 h. On average, ants removed 63.8% of myrmecochorous seeds and 10.9% of seeds without adaptation to ant dispersal. Analysis of soil seed content revealed that myrmecochores, in spite of expectations that they would accumulate in nests of seed dispersing ants, were most abundant in the soil of control plots. Evidence on seed relocation to the ant nests was obtained from a comparison of mounds of seed dispersing and seed non-dispersing ant species, as more seeds were found in the mounds of Formica spp. and Tetramorium caespitum (seed dispersers) in comparison with the mounds of Lasius favus (non-disperser).The soil seed bank of the compared microhabitats (control plots and mounds of 3 ant species) differed in their species composition, seed abundance and vertical distribution. The most distinct qualitative differences were between seed flora of control plots and mounds of Tetramorium caespitum. Control plots had approximately 30,000 propagules per m², which was double the number of seeds found in the ant mounds. In control plots, abundance and diversity of seeds steeply declined with depth; this trend was not observed in the mounds probably due to bioturbation. In the seed survival experiment, more seeds (2 out of 3 species) survived in control plots, which may also contribute to the higher seed abundance in this microhabitat.This study showed that seed relocation by ants does not contribute significantly to seed bank build-up at this study site. Ants may, however, increase the regeneration success of myrmecochores, mainly by dispersal for distance and placement in a larger spectrum of microsites, in contrast to species not adapted for myrmecochory.     

Interaction between the non-myrmecochorous herb Galium aparine and the ant Formica polyctena

Non-myrmecochorous plant species are able to colonize Formica polyctena mounds. Galium aparine has been chosen as a model to study this kind of ant-plant association. In order to understand benefits to ants and plants, we have studied removal rates of fruits by ant workers, and compared soil seed material, seedling pattern and development of plants of the nest versus forest.Ant workers remove fruits at very low rates; removal time for half of the sample of 10 seeds was on average 5 h. Removed fruits were usually lost within 20–30 cm distance of the ant route. The soil samples from the nests contained a higher number of fruits, than those from randomly chosen sites in the forest. Seeds start to germinate in late August; on the F. polyctena mounds significantly earlier than at surrounding places. The seedling density was higher within experimental quadrats located on mound sites especially in late September–early October. The highest density of seedlings was on the slope of the mound. Plants from mounds are significantly longer, have a higher number of nodes during all stages of development and a higher mortality rate than that at surrounding places.The interactions between G. aparine and F. polyctena ants are asymmetrical. In spite of higher mortality rate of seedlings on mounds, ants provide some benefits to the plant. Firstly, workers are additional dispersal agents for G. aparine fruits. Furthermore, the soil peculiarities of ant mounds are beneficial for seed germination and seedling growth. G. aparine does not provide ants with energetically valuable food like plants with elaiosome-bearing seeds. Moreover, the high density of seedlings and plants on the mounds shades the nest.     

热带森林不同演替阶段蚂蚁巢穴的分布特征及其影响因素

为探明热带森林蚂蚁巢穴的分布特征及其影响因素, 采用样方法研究了西双版纳不同演替阶段热带森林定居巢穴蚂蚁的种类及其巢穴的密度、盖度和空间分布特征, 并分析了土壤理化环境与蚂蚁种类总数、巢穴密度及盖度的相关性。结果表明, 不同演替阶段热带森林蚂蚁种类总数、巢穴的密度及盖度大小顺序为: 小果野芭蕉 (Musa acuminata)群落>白背桐(Mallotus paniculatus)群落>思茅崖豆(Mellettia leptobotrya)群落, 并且热带森林的演替类型显著影响蚂蚁种类总数及巢穴密度, 而对巢穴盖度的影响未达到显著水平; 蚂蚁种类总数、巢穴密度与土壤总有机碳和水解氮显著正相关, 与土壤容重和土壤含水率显著负相关, 但所选择的土壤理化指标与巢穴盖度的相关性均未达到显著水平; 蚂蚁巢穴的空间分布呈随机分布格局。我们的数据表明, 不同演替阶段热带森林所形成的植被类型及土壤环境状况共同影响定居的蚂蚁种类总数与筑巢密度。     

Large grazers modify effects of aboveground-belowground interactions on small-scale plant community composition

Aboveground and belowground organisms influence plant community composition by local interactions, and their scale of impact may vary from millimeters belowground to kilometers aboveground. However, it still poorly understood how large grazers that select their forage on large spatial scales interact with small-scale aboveground–belowground interactions on plant community heterogeneity. Here, we investigate how cattle (Bos taurus) modify the effects of interactions between yellow meadow ants (Lasius flavus) and European brown hares (Lepus europaeus) on the formation of small-scale heterogeneity in vegetation composition. In the absence of cattle, hares selectively foraged on ant mounds, while under combined grazing by hares and cattle, vertebrate grazing pressure was similar on and off mounds. Ant mounds that were grazed by only hares had a different plant community composition compared to their surroundings: the cover of the grazing-intolerant grass Elytrigia atherica was reduced on ant mounds, whereas the relative cover of the more grazing-tolerant and palatable grass Festuca rubra was enhanced. Combined grazing by hares and cattle, resulted in homogenization of plant community composition on and off ant mounds, with high overall cover of F. rubra. We conclude that hares can respond to local ant–soil–vegetation interactions, because they are small, selective herbivores that make their foraging decisions on a local scale. This results in small-scale plant patches on mounds of yellow meadow ants. In the presence of cattle, which are less selective aboveground herbivores, local plant community patterns triggered by small-scale aboveground–belowground interactions can disappear. Therefore, cattle modify the consequences of aboveground–belowground interactions for small-scale plant community composition.     

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     

Influence of small-scale disturbances by kangaroo rats on Chihuahuan Desert ants

Banner-tailed kangaroo rats (Dipodomys spectabilis) are prominent ecosystem engineers that build large mounds that influence the spatial structuring of fungi, plants, and some ground-dwelling animals. Ants are diverse and functionally important components of arid ecosystems; some species are also ecosystem engineers. We investigated the effects of patch disturbances created by D. spectabilis mounds on ant assemblages in a Chihuahuan Desert grassland in southern New Mexico by using pitfall traps in a paired design (mound vs. matrix). Although the disturbances did not alter species richness or harbor unique ant communities relative to the matrix, they did alter species composition; the abundances of 6 of 26 species were affected. The disturbances might also act to disrupt spatial patterning of ants caused by other environmental gradients. In contrast to previous investigations of larger-scale disturbances, we detected no effects of the disturbances on ants at the functional-group level. Whether ant communities respond to disturbance at a functional-group or within-functional-group level may depend on the size and intensity of the disturbance. Useful functional-group schemes also may be scale-dependent, however, or species may respond idiosyncratically. Interactions between disturbance-generating mammals and ants may produce a nested spatial structure of patches. Received: 11 October 1999 / Accepted: 11 March 2000     

Effects of ants on plant diversity in semi-natural grasslands

Ants (L. niger and L. flavus) build conspicuous mounds that are covered with vegetation. The aim of this study was to investigate whether the vegetation on ant mounds in semi-natural grasslands differed from that around the mounds. Another aim was to investigate whether the changes in the vegetation on ant mounds were influenced by grazing management or by habitat characteristics, semi-dry versus moist. Here, the total number of plant species and total plant cover were lower on ant mounds than in patches off-mound. The plant cover of perennials that form rosettes was twice as high on mounds inhabited by L. niger than on those inhabited by L. flavus. Only a few plant species were restricted to either ant mounds or adjacent field and the effects of ants on the plant diversity in semi-natural grasslands seemed to be low. Grazing management did not affect the differences in the vegetation on ant mounds and in equal-sized patches off-mound, whereas habitat characteristics affected ant-induced changes in vegetation cover of some plant species.     

Influence of mound construction by red and hybrid imported fire ants on soil chemical properties and turfgrass in a sod production agroecosystem

Mound construction by imported fire ants (Solenopsis spp.) actively modify the biogeochemical and physical properties of soil; however, their influence on nutrient levels in surrounding vegetation is poorly understood. Aside from the reported persistence of elevated available P and K levels in clay-rich soils one year after mound abandonment, the relative stability of nutrient concentrations from one season to the next is largely unknown. Nutrient concentrations were concurrently analysed from ant mounds and undisturbed soils as well as plant samples collected from warm-season turfgrass in a commercial sod production agroecosystem. Initial collection of soil and turfgrass samples coincided with peak annual biomass (September 2006); the second soil sample collection occurred over twelve weeks later during turfgrass dormancy and ant brood minimum (December 2006). Total C, C/N ratios, organic matter (OM), and Zn²⁺ concentrations as well as pH of ant mound soils were significantly higher than control plot soils; these trends persisted across seasons. Turfgrass harvested from ant mound perimeters in September exhibited elevated N, P, Ca²⁺, S, Cu²⁺, Fe²⁺, and Na⁺ concentrations. Evaluation of the relative stability of soil parameters across seasons revealed a significant drop in ant nest pH from September to December 2006. Total N of mound soils was distinctively greater than control soil counterparts during September only. Soil P, K⁺, Ca²⁺, Mg²⁺ and S (all macronutrients), as well as Na⁺ concentrations from ant mound soils were substantially elevated during the late Autumn to Winter transition compared to control soil locations, whereas Fe²⁺ and Mn²⁺ levels (both micronutrients) were significantly lower in ant mound soils versus control soil environments. Continuous pedoturbation by imported fire ants as well as seasonal shifts in mound soil chemistry resulting from changes in assimilation/dissimilation among mound biota may influence the site-specific effectiveness of microfaunal pathogens (e.g., Thelohania solenopsae) or parasites (e.g., Orasema spp.) identified as classical biological control agents of non-native Solenopsis spp. Therefore, further study of the intrinsic complexities of soil ecosystem dynamics of imported fire ant mounds across several seasons is warranted. Received 10 September 2007; revised 23 April 2008; accepted 1 May 2008.     

Interactions between above- and belowground biota: importance for small-scale vegetation mosaics in a grassland ecosystem

Grasslands are often characterised by small-scale mosaics in plant community composition that contribute to their diversity. Although above- and belowground biota can both cause such mosaics, few studies have addressed their interacting effects. We studied multi-trophic interactions between aboveground vertebrate grazers, subterranean ants, plant-pathogenic soil biota (especially nematodes) and the vegetation in a temperate grassland. We found that when rabbits and cattle locally omit vegetation patches, yellow ants ( Lasius flavus ) respond to the taller vegetation by digging up more sand from deeper soil layers (hence making taller nest mounds), probably to maintain sufficiently high soil temperatures. We found that this ant digging affects other soil biota, as the mounds contain fewer plant-parasitic and fungivore nematodes. Also, the mounds have lower moisture content and soil bulk densities, and higher pH and available nutrient content than the directly surrounding soil. The clonal sedge Carex arenaria grows vigorously on the mounds, producing more shoots and shorter rhizome internode lengths than in surrounding vegetation. Other plant species, such as the grass Festuca rubra , dominate the surrounding vegetation. A greenhouse bioassay experiment revealed that harmful soil organisms (as plant-parasitic nematodes and pathogenic fungi) outweighed the effect of beneficial organisms (e.g., mycorrhizae) in this system. Rhizome biomass and shoot production of C. arenaria were indeed inhibited less by biota in soil from ant mounds than by biota in soil from the surrounding vegetation . However, the total biomass production of C. arenaria was inhibited as strongly in both soil types. F. rubra was inhibited more strongly by biota in the surrounding soil. We suggest that various direct and indirect interactions between above- and belowground biota can contribute to community mosaics and hence diversity in grasslands.     

The use of the mounds of Lasius fiavus in teaching some principles of ecological investigation

The mounds built by the yellow hill ant, Lasius fiavus, in British pastures have striking and consistent effects on the patterns of plant species. Winter annuals, for example, are often confined to the mounds, but many rosette perennials, though abundant in the surrounding grassland, are virtually absent from ant-hills. A comparison of the abundances of six or seven species on and off the mounds provides a field practical suitable for senior school students and undergraduates. The students discuss the sampling methods to be used, collect the data, analyse it, and discuss the factors which might influence the relative abundance of the plant species. This paper describes the practical in detail, and provides some background information about the ecology of the ants and some common grassland plants. Some suggestions for individual projects are put forward.     

Spatial distribution of ant mounds and effects on soil physical properties in wetlands of the Sanjiang plain, China

Ants constitute a dominant element of soil mesofauna due to their biomass, abundance, richness of species and distribution within terrestrial ecosystems. They are important regulators of soil aggregate structure as they translocate large amounts of soil from the bottom to the soil surface. In doing so, they form biogenic structures (BS) made up of aggregates of different sizes and characteristics, i.e. ant mounds. These BS have varying characteristics according to the ant species and the soil where they carry their activities. Ants are considered soil engineers because of their effects on soil properties, availability of resource and flow of energy and nutrients in soil. Thus, it is important to gain information on their distribution and abundance. Relatively little is known about the spatial distribution of mounds and their role in the soil physical properties in wetlands of the Sanjiang plain, China. We conducted a survey of ant mounds and measured the density, height, and diameter and material composition of different ant mounds. The ecological characteristics of wetlands that ant mounds wide occurrence were also investigated, including soil type, hydrology characters and plant composition. Differences in soil particle composition, bulk density and soil moisture between ant mound and natural meadow were measured to assess the influences of ant mounds on soil physical properties. We also studied the effects of ant mounds on the microtopography of meadows. Ant mounds were found mainly in the transition zone between terrestrial and aquatic habitats, with wetland type, including Calamagrostis augustifolia wet meadow, C.augustifolia marsh meadow, shrubs marsh meadow and Carexmeyeriana–Carexappendiculata wetland, being a significant factor. Most of the mounds detected were inhabited by Lasius flavus Fabricius, Lasius niger Linnaeus and Formica sanguinea Latreille, which occupied 52.9%, 26.5% and 20.6% of the mounds surveyed, respectively. The density, height, diameter and mound composition were significantly different among the mounds of F. sanguinea Latreille, L. flavus Fabricius and L. niger Linnaeus. The average density and diameter of L. flavus mounds was significantly higher than those of other ant mounds. The average height of F. sanguinea mounds was highest among the mounds detected. Mound building activities changed soil particle size distribution, with the silt and clay content of mounds higher than for non-mound soil. Compared with adjacent, non-mound soil, the bulk density (0–30 cm) and water content (0–25 cm) of mound soil were significantly lower, but there were no significant differences between the mound soil of F. sanguinea Latreille and L. flavus Fabricius. The spatial distribution of ant mounds with different height and diameter also changed the micro-geomorphology of the soil surface, increasing the degree of fluctuation of the microtopography. The ant distribution characteristics and their ecological roles respond to a wide range of environmental alterations. The biogenic structures of ant and the specific environment associated with them have been defined as the “functional domain”, a sphere of influence that may significantly affect soil processes at certain spatial and temporal scales. Our results suggest that the distribution and structure of ant mounds can indicate wetland environmental changes, with mounds influencing ecosystem functions and enhancing wetland degradation.     

Testing associational resistance against pine weevils mediated by Lasius ants attending conifer seedlings

Natural enemies attracted to plants may provide those plants with protection against herbivores but may also protect neighbouring plants, that is through associational resistance. Ant attendance may be an important mechanism for the occurrence of such effects because ants can reduce the damage caused by a wide variety of herbivorous insects. Ants have been shown, in a previous field experiment, to decrease the damage caused by the pine weevil, Hylobius abietis (L.) (Coleoptera: Curculionidae), a pest species that causes high seedling mortality in forest regeneration areas. In this study, we specifically tested whether seedlings planted close to ant‐attended seedlings experience associational resistance. We did this under laboratory conditions using the ant species Lasius niger (L.) (Hymenoptera: Formicidae). The feeding damage by pine weevils was significantly reduced on seedlings attended by ants. The neighbouring seedlings, however, did not experience associational resistance. Nevertheless, some associational effects were observed as the number of weevils recorded on both ant‐attended and neighbouring seedlings was significantly lower compared with ant‐excluded seedlings.     

Vegetation heterogeneity caused by an ecosystem engineer drives oviposition-site selection of a threatened grassland insect

Soil-disturbing ecosystem engineers play an important role in plant-species diversity in grasslands as they increase vegetation heterogeneity by creating gaps due to burrowing or mound-building activities. However, knowledge of the ecological importance of these microsites for arthropods is still rare. In this study, we analyse the role of ant-nest mounds of the yellow meadow ant (Lasius flavus) for oviposition-site selection of the silver-spotted skipper (Hesperia comma). Ant mounds were searched for H. comma eggs. Microclimatic and vegetation parameters were ascertained at occupied sites and control sites within the matrix vegetation. Furthermore, we analysed the habitat requirements of L. flavus by means of nest counting and the sampling of environmental parameters within different sites. L. flavus occurred most frequently in abandoned and less steep sites with deeper soils. Mean egg occupancy rates of H. comma on ant hills were 32 %, nearly twice as high as at control sites (18 %). In contrast to the surrounding vegetation, nest mounds were characterized by a lower vegetation cover and litter and a higher proportion of bare ground. Furthermore, they had a higher cover of host plants compared with control samples. These microhabitats offered the following essential key factors for the larval development of H. comma: (1) a suitable microclimate due to open vegetation and (2) a high amount of host plants. This study highlights the importance of L. flavus as an ecosystem engineer within central European grasslands because this species increases vegetation heterogeneity.