Ant-induced soil modification and its effect on plant below-ground biomass

Lasius flavus is a dominant mound-building ant species of temperate grasslands that significantly modifies soil parameters. These modifications are usually the result of workers’ activities such as food accumulation and nest construction. An alternative hypothesis that could explain changes in soil is colony founding in areas of higher soil fertility.In our study we investigated several soil parameters sampled in 10 ant nests and adjacent (control) plots in mountain grassland in Slovakia. The alternative hypothesis was tested by comparing occupied and abandoned mounds. While we found increased concentrations of available P and K in the nests, concentrations of total C, total N, Ca²⁺ and Mg²⁺ were lower there. We propose that differences found between the soil of nests and control plots are entirely a product of ant activity during mound occupancy and not due to initial soil differences during nest establishment. This was confirmed by the comparison of occupied and abandoned nests in which the soil fertility of abandoned nests was similar to conditions in the surrounding soil.Along with the modification of soil chemistry, we recorded changes in soil physical properties and the vertical distribution of nutrients. Ant nests were characterized by the dominance of 0.02–0.1 mm particles and lower bulk density. In the same habitat, nutrient concentrations did not change along the vertical gradient in contrast to control plots where soil nutrients decreased and bulk density increased with depth. Root biomass followed the vertical pattern observed with nutrients: in control plots, most roots were concentrated in the uppermost layer (0–3 cm), whereas they were evenly distributed along the vertical gradient in the nests. We also found that rhizome internodes of Agrostis capillaris were thinner and longer in plants from the mounds. Changes in soil physical properties, vertical distribution of nutrients and root biomass in the nests are most probably a consequence of mounding and soil mixing (bioturbation), which has been less reported on in ant-soil studies.     

Some relations of the meat ant, Iridomyrmex purpureus (hymenoptera: formicidae) with soil in South Australia

Iridomyrmex purpureus F. Smith is common in southern Australia and nests in conspicuous. often gravel-covered mounds. The effects of the ant on some properties of soil in its nests, and the effects of soil type on the distribution of the ant were studied by comparing the distribution of particle sizes in soil of nests and undisturbed soil profiles. Particle size analysis was used also to determine the relationships of nests and more or less truncated profiles on a naturally eroding, lateritic podzolic soil. Nests are constructed by excavation of gallery systems in the upper part of the profile, with a few shafts extending downwards for 1 metre or more. Excavated material is incorporated in the mound which is usually composed of soil from the A horizon with additions from the B horizon and imported gravel. The density and rate of turnover of nests both appear to he too low for I. purpureus to have any significance to pedogenesis. But soil texture and consistency do influence the distribution of the ant. for nests are not constructed in sands or in very loose, structureless soils. Examination of the distribution of particle sizes in material from nests on the lateritic podzolic soil suggests that establishment of some existing mounds may have pre-dated erosion which has, subsequently, removed more than 20 cm of A horizon. The erosion process would probably require at least hundreds of years, and. if the interpretation of these nests is correct, suggests that they are very old. This is not inconsistent with other observations on rates of change in this population of I. purpurens.     

Unique soil microbial assemblages associated with grassland ant species with different nesting and foraging strategies

Ants are important ecosystem engineers and can be abundant in extensively managed grassland ecosystems. Different ant species create nests varying in structure and size, and tend to have a variety of feeding strategies. Differences in food imported to the nest and contrasting nesting behaviour may control soil microbial community structure in nest soil, with cascading effects on nutrient cycling, but this has not been tested in grassland ants. Soil and ants were sampled from nests of three ant species: two formicines; Lasius flavus (aphid farmer/scavenger, mound builder) and Formica lemani (scavenger/hunter, non-mound builder), and a myrmicine; Myrmica sabuleti (hunter/scavenger, non-mound builder), in an extensively grazed temperate grassland and compared to similar soils without ants. Microbial assemblages were determined using molecular approaches (terminal restriction length polymorphism and automated ribosomal intergenic spacer analysis). Both aboveground (vegetation diversity) and belowground (soil physico-chemical properties) components were measured to assess the potential of the different ant species to modify the environment. Stable isotope ratios (δ¹³C and δ¹⁵N) of ant tissues and nest soil organic matter confirmed differences in trophic distances. Significant changes in soil pH, moisture content, total C and total N, and in vegetation composition, demonstrated ant ecosystem engineering effects. In turn, nests of L. flavus, M. sabuleti and F. lemani had different microbial activities and harboured significantly different microbial assemblages (total bacteria, total fungi, ammonia-oxidising bacteria and nitrogen-fixing bacteria), but the diversity was similar. These findings suggest that grassland ants can control microbial assemblages via changes in physical and biological soil characteristics in their nests, and as such, different ant species harbour unique microbial assemblages in nests.     

The influence of different vegetation patches on the spatial distribution of nests and the epigeic activity of ants (Lasius niger) on a spoil dump after brown coal mining (Czech Republic)

A study was carried out during 2001 on mine tailings in NW Bohemia aimed at describing the spatial patterns of nests distribution and epigeic activity of ants in relation to the vegetation mosaic. Lasius niger was the most abundant species of ant and its nest mounds were significantly more numerous in patches with sparse vegetation than inside dense Calamagrostis epigejos vegetation; this was particularly true for small and medium-sized nests. Small and medium nests also occurred more frequently near the edges of a given patch than in the center. Large and medium nests were randomly distributed in the area, whereas small nests had an aggregated distribution. Pitfall trapping reveal significantly higher activity of L. niger workers in tall and dense vegetation stands in comparison with low and sparse vegetation. This pattern was particularly pronounced during the peak of foraging activity in summer and was not so significant in spring or autumn. We expect that ant preferentially forage in shaded habitats during the summer months when bare soil may be too hot. The results indicated that nesting and foraging may differ in their microclimatic requirements and the formation of vegetation mosaics may be important to changes in the ant population during succession.     

Impact of subterranean fungus-growing termites (Isoptera, Macrotermitiane) on chosen soil properties in a West African savanna

Fungus-growing termites (Isoptera, Macrotermitinae) play an important role in tropical ecosystems in modifying soil physical properties. Most of the literature regarding the impact of termites on soil properties refers to termite epigeous mounds. In spite of their abundance and activity in African savannas, few studies deal with the properties of underground nest structures (fungus-comb chambers) built by subterranean Macrotermitinae termites. We tested whether these termites significantly modify the soil physico-chemical properties within their nests in a humid tropical savanna and whether these effects are different for two termite species with differing building behaviour. Termite-worked soil material was collected from fungus-comb chamber walls of two widespread species: Ancistrotermes cavithorax, which builds diffuse and ephemeral nests and Odontotermes nr pauperans, which most often builds concentrated and permanent nests for a comparatively much longer period of time. Neither species influenced soil pH but both significantly modified soil texture and C-N content in their nest structures. A strong impact on clay-particle size was also detected but no significant differences in clay mineralogy. Thus Odontotermes has a greater effect on soil properties, that could be explained by its building behaviour and the concentration in space of its nest units. Therefore, spatial pattern and life-span of fungus-comb chambers should be an important parameter to be considered in the functional role of subterranean Macrotermitinae termites in the savanna.     

Physical, chemical and phosphatase activities characteristics in soil-feeding termite nests and tropical rainforest soils

Little is known about the relationship between soil biological function and the physical and chemical characteristics of soil-feeding termite nests in the Lopé tropical rainforest (Gabon). We compared nine soil-feeding termite nests of Cubitermes of different ages (fresh to mature to old) and six surrounding soils that originated from three forests differing with respect to age and vegetative cover according to 14 physical and chemical variables and acid (pH 4) and alkaline (pH 9) phosphatase activities. Physical and chemical variables of the studied samples were influenced by the three factors tested: (1) forest age, (2) termite activity (nest versus soil), (3) termite nest age. Soils from the gallery forest were strongly discriminated from all the other soils studied notably due to their high organic matter contents. All mature nests showed significant increases in K, P, clay and fine silt, pH, and cationic exchange capacity compared to soils. Some nests also had increased amounts of organic matter and larger water retention capacities. Moreover, we observed that with age the termite nests possessed decreased values of these variables from fresh to mature to old. Likewise, phosphatase activities also differed according to the three factors tested. Due to its high organic matter contents, the highest phosphatase activities were noted in the gallery forest. Within each forest, phosphatase activities decreased in mature nests compared to soils and tended to be higher in fresh nests compared to mature nests. These differences might be due to an inhibition by high inorganic P contents, as mature nests were enriched in this element and to the quality of organic matter as nests are built with termite faeces. Termite activity has an important role in influencing physical and chemical variables and phosphatase activities.     

Use of RAPD markers for the study of microbial community similarity from termite mounds and tropical soils

In this study, we test the use of the RAPD (Random Amplified Polymorphic DNA) molecular markers as a way to estimate the similarity of the microbial communities in various termite mounds and soils. In tropical ecosystems, termite activities induce changes in the chemical and physical properties of soil. The question then arises as to whether or not termites affect the presence of natural microbial communities. Successful 16S rDNA amplifications provided evidence of the occurrence of bacterial DNA in termite constructions including both soil feeder and fungus grower materials. A phenetic dendrogram using the similarity distance calculated from pairwise data including 88 polymorphic RAPD markers was reconstructed and bootstrap scores mapped. The microbial communities of the mounds of the four soil-feeding termites were clustered in the same clade, while those of the mounds of the fungus-growing species were distinct like those of control soils. Microbial changes in nests result from termite building behavior, depending on whether they include feces in their constructions for soil-feeders or use saliva as particle cement for fungus-growers. It is argued that RAPDs are useful markers to detect differences in microbial community structure not only between termitaries and control soils but also between mounds of soil-feeders.     

Biological Sources of Polycyclic Aromatic Hydrocarbons (PAHs) in the Amazonian Rain Forest

Hazardous polycyclic aromatic hydrocarbons (PAHs) widely occur in the environment and are believed to be mainly anthropogenic. Here we present strong indications for large biological sources of the PAHs naphthalene, perylene, and possibly also phenanthrene in the Amazonian basin. Termite nests, plant wood, and soils were sampled. Naphthalene is detected in plant wood and is accumulated in the nests of termites from the genus Nasutitermes. Perylene is found in all studied termite nests including six different genera. Phenanthrene occurs at substantial concentrations in wood, soil, and termite nests.     

The impact of harvester ants on decomposition, N mineralization, litter quality, and the availability of N to plants in the Mojave Desert

In arid areas of North America, nests of the seed-harvesting ant Pogonomyrmex rugosus tend to be elevated in mineral nitrogen and other soil nutrients relative to other microhabitats. We investigated the roles of decomposition, N mineralization, and plant nutrient uptake in maintaining high standing stocks of nutrients in P. rugosus ant nests. Decomposition rates of standard cellulose substrates placed on the surface of ant nests and other desert microhabitats suggest that conditions found in ant nests and bare areas are conducive to higher rates of decomposition than conditions under shrubs. In laboratory incubations of moist soil, net N mineralization rates were significantly higher in soil from ant nests than from bare areas and under two of three plant species. Net N mineralization rates measured in situ were much lower than those measured in laboratory incubations, but ant nest soil still exhibited higher rates at one of two sites. Litter collected from ant mounds, composed chiefly of seed chaff, was similar in N content to litter collected from underneath the dominant plant species, but had a significantly higher mean δ¹⁵N. Using this distinctive isotope signature as a tracer, we found no evidence that large perennial shrubs tap ant nests as a source of N. An invasive, annual grass species was significantly enriched in ¹⁵N, had higher leaf %N, and produced more seeds when growing on the mound than when growing several meters away; however P. rugosus nest surfaces are typically free of such annuals. We conclude that both high rates of nutrient cycling relative to other Mojave Desert microhabitats and low N utilization by the surrounding vegetation contribute to high standing stocks of mineral N in P. rugosus nests.     

Ants mediate soil water in arid desert ecosystems: Mitigating rainfall interception induced by biological soil crusts?

As vital components of desert systems, the roles of ants in arid ecological processes have been well documented, while little attention has been given to their effects on soil water. We conducted a six-year investigation in sand dune systems stabilized via revegetation, to explore the hydrological role of ants through comparing the influence of ant nests on rainfall infiltration in different-aged revegetated dunes. The presence of ant nests markedly enhanced infiltration due to weakening the rainfall interception by biological soil crusts (BSCs) in revegetated dunes. The distribution of ant nest was denser in older revegetated areas, due to better developed BSCs of later successional stages, compared to younger revegetated areas. Ants prefer later to early successional BSCs because the later lichen–moss dominated crusts were thicker and their surface was more stable than the early cyanobacteria dominated crusts. Conversely, the crustal rainfall interception was positively correlated with BSC thickness. These findings suggest that the occurrence of ant nests in older revegetated areas benefited to the planted shrubs with deeper root systems and maintain a relative constant cover of shrubs in artificial sand-binding vegetation following an increase in infiltration to deeper soil layers.     

Temporal patterns of nutrient availability around nests of leaf-cutting ants (Atta colombica) in secondary moist tropical forest

Leaf-cutting ants consume up to 10% of canopy leaves in the foraging area of their colony and therefore represent a key perturbation in the nutrient cycle of tropical forests. We used a chronosequence of nest sites on Barro Colorado Island, Panama, to assess the influence of leaf-cutting ants (Atta colombica) on nutrient availability in a neotropical rainforest. Twelve nest sites were sampled, including active nests, recently abandoned nests (<1 year) and long-abandoned nests (>1 year). Waste material discarded by the ants down-slope from the nests contained large concentrations of nitrogen and phosphorus in both total and soluble forms, but decomposed within one year after the nests were abandoned. Despite this, soil under the waste material contained high concentrations of nitrate and ammonium that persisted after the disappearance of the waste, although soluble phosphate returned to background concentrations within one year of nest abandonment. Fine roots were more abundant in soil under waste than control soils up to one year after nest abandonment, but were not significantly different for older sites. In contrast to the waste dumps, soil above the underground nest chambers consistently contained lower nutrient concentrations than control soils, although this was not statistically significant. We conclude that the ‘islands of fertility’ created by leaf-cutting ants provide a nutritional benefit to nearby plants for less than one year after nest abandonment in the moist tropical environment of Barro Colorado Island.     

Leaf-cutting ant (Atta Sexdens) and nutrient cycling: deep soil inorganic nitrogen stocks, mineralization, and nitrification in Eastern Amazonia

Nest excavation and agricultural activities of the leaf-cutting ant Atta sexdens create complex belowground heterogeneity in secondary forests of Eastern Amazonia. We examined the effects of this heterogeneity on inorganic-N stocks, net mineralization, and net nitrification to test the hypothesis that the bulk soil of the nests has higher net rates of mineralization and nitrification than soil that was not affected by the influences of ant nests, throughout the profile. This study was conducted in a secondary forest at Fazenda Vitoria, near Paragominas in the Eastern Brazilian Amazon, where a previous study showed that the bulk soil of ant nests had elevated NO₃⁻. The results of the inorganic-N measurements were consistent with the previous study, showing elevated NO₃⁻ deep in the soil profile of the nests. However, neither net mineralization nor net nitrification were significantly greater at depth in the mineral soil of the nests compared to soil that was not influenced by nests (P=0.05), although variability was higher in the nest soil. These results suggest that the NO₃⁻ may have diffused into the surrounding mineral from the N-rich organic matter buried by the ants in chambers within the deep soil.     

Increased CO2 emission and organic matter decomposition by leaf-cutting ant nests in a coastal environment

Leaf-cutting ants perform a vital role in the cycling of carbon and nutrients in tropical ecosystems. Nests have high levels of organic matter and refuse dumps host up to two times more soil micro-organisms than non-nest soil. The increased levels of organic matter in the soil of nests, however, can affect CO₂ emissions from soil and alter the balance of atmospheric CO₂. We aimed at assessing the effect of nests of the leaf-cutting ant Acromyrmex balzani on CO₂ emissions in a coastal area of Northeast Brazil. Results show that A. balzani nests emitted up to four times more CO₂ than the surrounding soil and emissions were positively correlated with soil moisture and soil organic matter (SOM) content. In addition, field experiments demonstrated that refuse material has a lower residence time than the leaf material brought to the colonies. Despite the high density of nests and high content of SOM compared to adjacent control soil, CO₂ emissions by A. balzani nests represent only 0.3% of the total CO₂ efflux by the studied ecosystem. Although these effluxes account for a relative small portion of the total soil CO₂ emission, they are still important for the understanding of C balance, especially when one considers the thousands of tons of CO₂ emitted each day, across entire Neotropical regions where leaf-cutting ants occur.     

Nest-mounds of the yellow meadow ant (Lasius flavus) at the “Alter Gleisberg”, Central Germany: Hot or cold spots in nutrient cycling?

Nests of the yellow meadow ant (Lasius flavus) occur at high densities in grasslands worldwide. Although many studies have shown that L. flavus nests influence soil nutrient contents, little is known about their effect on soil nutrient cycling rates. The aim of this study was to examine the role of nest-mounds inhabited by L. flavus as potential ‘hot spots’ for soil nutrient cycling. Six pairs of nest-mounds and control soils were selected at a grassland site at the plateau of the Alter Gleisberg (Thuringia, Central Germany). L. flavus significantly modified the soil environment within the nest. In comparison to the control soils, nest-mounds were characterized by slightly higher soil temperatures during the summer months. In addition, we found that nests were related to decreased potential C mineralization rates and increased potential net N mineralization rates. Nest-mound soil exhibited lower amounts of SOC, hot-water extractable DOC and DN, and higher concentrations of leachable DOC and DN. Moreover, ants promoted the enrichment of base cations in the nest. Differences in the soil environment between nests and control soils were possibly a result of the burrowing activity of ants, soil mixing, accumulation of aphid honeydew, and decreased plant-derived nutrient inputs into the nest-mound soil. In conclusion, L. flavus nest-mounds had a significant but element dependent effect on the soil nutrient cycling and may represent cold spots for C cycling and hot spots for N cycling. Thus, L. flavus nests increase the spatial heterogeneity of soil properties and create unique micro-sites within grassland ecosystems.     

Biological activity of ant nests in the middle taiga zone

Ants are the most widespread social insects. Many species of ants influence a range of soil properties when building their nests in soil. The paper discusses the effects of ants on soil biological activity (nitrogen fixation, denitrification), as well as on biomass and the diversity of microorganisms. In many cases, these processes were more active in anthills. Changes in soil pH caused by nest-building activity of ants were also. There were no patterns in the distribution of pH values. Specific differences between microbial complexes of anthills and corresponding reference soils were revealed.     

Do long-lived ants affect soil microbial communities?

This study was designed to test the hypothesis that desert ant species that build nests that remain viable at a particular point in space for more than a decade produce soil conditions that enhance microbial biomass and functional diversity. We studied the effects of a seed-harvester ant, Pogonomyrmex rugosus, and two generalist ant species, Aphaenogaster cockerelli and Myrmecocystus depilis, on soil microbial communities. Microbial biomass was higher in P. rugosus-modified soils than in reference soils when soil water content was higher than 3%. Microbial biomass was either higher in reference soils or exhibited no difference in reference soils and nest-modified soils of A. cockerelli and M. depilis. There were differences in microbial functional diversity and microbial community level physiological profiles (MicroResp method) between ant-nest-modified and reference soils of the three ant species on some sampling dates. Temporal patterns of soil microbial communities associated with the ant species resulted from differences in soil moisture, density, and species composition of the annual plant communities associated with the ant nests and in reference areas. Differences in annual plant communities associated with ant nests and surrounding areas resulted in different chemical inputs into the soil organic-matter pools. This study shows that generalizations about the effects of long-lived ant nests on soil biota in arid regions must consider feeding behaviors of the ant species and temporal patterns of rainfall.     

Effectiveness of different conservation measures for loggerhead sea turtle (Caretta caretta) nests at Zakynthos Island, Greece

In this paper, we evaluate the effectiveness of different well-tested conservation techniques in an effort to successfully protect sea turtle nests. From an eight-year study on the island of Zakynthos, West Greece, we have experimentally investigated the effectiveness of two different conservation techniques applied to loggerhead sea turtle nests and provided statistical measures to evaluate their conservation value. The categories of nests evaluated include: (i) nests incubated in situ, (ii) translocated into a beach hatchery, or (iii) protected by metal cages. Results of the analysis showed significant interannual variations in hatching success as obtained for each one of the three groups of nests. Significant differences were also observed when comparing hatching success data among the three groups of nests during the eight-year period. Overall, our results indicate that relocation of nests laid at highly threatened locations and the placement of protective cages on nests in situ provide adequate conservation measures that could allow an increase in hatchling production; although their choice and application should be based on the specific conditions and threats of each nest.     

Phosphorus forms as affected by abandoned anthills (Formica polyctena Förster) in forest soils: sequential extraction and liquid-state 31P-NMR spectroscopy

Ants are known to concentrate phosphorus (P) inside their nests via collection of food and litter. To elucidate the possible effects on long-term availability of soil P, five anthills abandoned by Red wood ant (Formica polyctena Förster) > 5—20 years ago were characterized for soil P forms in a temperate Danish deciduous forest. Sequentially extracted P fractions and liquid-state ³¹P nuclear magnetic resonance (NMR) spectra were obtained on surface samples (0—10 cm) from abandoned anthills and adjacent topsoil; in addition one representative soil profile in an abandoned anthill was investigated. The results show that different inorganic and all organic P fractions were enriched by a factor of 2.0—3.3 inside anthills relative to the surrounding soil. The soil underneath the abandoned anthill had higher P contents until 50-cm depth. Phosphorus composition was less affected by former anthill construction. Only the younger anthills revealed a preferential accumulation of labile organic P forms such as Na-HCO₃ extractable P or diester P. The accumulation of the stable and moderate labile P forms, however, persisted for ≥ 20 years after abandonment. We concluded that former ant activity enhanced long-term P availability of soil due to high local P inputs, whereas changes of the P form distribution lasted 5—10 years after nest abandonment.     

Variability in soil chemistry and arbuscular mycorrhizal fungi in harvester ant nests: the influence of topography, grazing and region

Harvester ants are important disturbance agents across western North America, but the effects of ant disturbances on soils may vary considerably with topography and land use. We examined how soil properties and arbuscular mycorrhizal (AM) fungi in harvester ant nests varied across spatial scales according to topography, grazing regime and region. Soils from undisturbed areas were compared with nest disturbances created by two species of harvester ants, Pogonomyrmex occidentalis on shortgrass steppe in Colorado and P. rugosus on Chihuahuan desert grassland in New Mexico, in 1996 and 1997. Nests of both ant species were enriched with NO₃^--N, total P and roots colonized by AM fungi. Soil moisture was higher in P. rugosus' nests and lower in P. occidentalis' nests compared to surrounding areas. Soil pH was consistently lower in ant nests. Broad-scale factors such as grazing, topography and site affected most soil properties in and away from ant nests. Site exerted a strong influence on soil organic matter, pH and moisture. Within sites, topography had a significant affect on pH. Mycorrhizal colonization was influenced by site and topography in 1996 only, a substantially wetter year at both sites. Lastly, nutrient levels were largely determined by the fine-scale effects of ant disturbances. Principal components analysis revealed that, after removing site-level effects, harvester ants have similar functional roles in creating soil heterogeneity in these two different semiarid ecosystems.