Impact of Biogenic Structures of the Soil-Nesting Ants Lasius niger and Lasius flavus on the Soil Microarthropod Community in Urban Green Spaces

Impact of Biogenic Structures of the Soil-Nesting Ants Lasius niger and Lasius flavus on the Soil Microarthropod Community in Urban Green Spaces | bioRxiv /* */ /* */ <!-- <!-- /*! * yepnope1.5.4 * (c) WTFPL, GPLv2 */ (function(a,b,c){function d(a){return"[object Function]"==o.call(a)}function e(a){return"string"==typeof a}function f(){}function g(a){return!a||"loaded"==a||"complete"==a||"uninitialized"==a}function h(){var a=p.shift();q=1,a?a.t?m(function(){("c"==a.t?B.injectCss:B.injectJs)(a.s,0,a.a,a.x,a.e,1)},0):(a(),h()):q=0}function i(a,c,d,e,f,i,j){function k(b){if(!o&&g(l.readyState)&&(u.r=o=1,!q&&h(),l.onload=l.onreadystatechange=null,b)){"img"!=a&&m(function(){t.removeChild(l)},50);for(var d in y[c])y[c].hasOwnProperty(d)&&y[c][d].onload()}}var j=j||B.errorTimeout,l=b.createElement(a),o=0,r=0,u={t:d,s:c,e:f,a:i,x:j};1===y[c]&&(r=1,y[c]=[]),"object"==a?l.data=c:(l.src=c,l.type=a),l.width=l.height="0",l.onerror=l.onload=l.onreadystatechange=function(){k.call(this,r)},p.splice(e,0,u),"img"!=a&&(r||2===y[c]?(t.insertBefore(l,s?null:n),m(k,j)):y[c].push(l))}function j(a,b,c,d,f){return q=0,b=b||"j",e(a)?i("c"==b?v:u,a,b,this.i++,c,d,f):(p.splice(this.i++,0,a),1==p.length&&h()),this}function k(){var a=B;return a.loader={load:j,i:0},a}var l=b.documentElement,m=a.setTimeout,n=b.getElementsByTagName("script")[0],o={}.toString,p=[],q=0,r="MozAppearance"in l.style,s=r&&!!b.createRange().compareNode,t=s?l:n.parentNode,l=a.opera&&"[object Opera]"==o.call(a.opera),l=!!b.attachEvent&&!l,u=r?"object":l?"script":"img",v=l?"script":u,w=Array.isArray||function(a){return"[object Array]"==o.call(a)},x=[],y={},z={timeout:function(a,b){return b.length&&(a.timeout=b[0]),a}},A,B;B=function(a){function b(a){var a=a.split("!"),b=x.length,c=a.pop(),d=a.length,c={url:c,origUrl:c,prefixes:a},e,f,g;for(f=0;f<d;f++)g=a[f].split("="),(e=z[g.shift()])&&(c=e(c,g));for(f=0;f<b;f++)c=x[f](c);return c}function g(a,e,f,g,h){var i=b(a),j=i.autoCallback;i.url.split(".").pop().split("?").shift(),i.bypass||(e&&(e=d(e)?e:e[a]||e[g]||e[a.split("/").pop().split("?")[0]]),i.instead?i.instead(a,e,f,g,h):(y[i.url]?i.noexec=!0:y[i.url]=1,f.load(i.url,i.forceCSS||!i.forceJS&&"css"==i.url.split(".").pop().split("?").shift()?"c":c,i.noexec,i.attrs,i.timeout),(d(e)||d(j))&&f.load(function(){k(),e&&e(i.origUrl,h,g),j&&j(i.origUrl,h,g),y[i.url]=2})))}function h(a,b){function c(a,c){if(a){if(e(a))c||(j=function(){var a=[].slice.call(arguments);k.apply(this,a),l()}),g(a,j,b,0,h);else if(Object(a)===a)for(n in m=function(){var b=0,c;for(c in a)a.hasOwnProperty(c)&&b++;return b}(),a)a.hasOwnProperty(n)&&(!c&&!--m&&(d(j)?j=function(){var a=[].slice.call(arguments);k.apply(this,a),l()}:j[n]=function(a){return function(){var b=[].slice.call(arguments);a&&a.apply(this,b),l()}}(k[n])),g(a[n],j,b,n,h))}else!c&&l()}var h=!!a.test,i=a.load||a.both,j=a.callback||f,k=j,l=a.complete||f,m,n;c(h?a.yep:a.nope,!!i),i&&c(i)}var i,j,l=this.yepnope.loader;if(e(a))g(a,0,l,0);else if(w(a))for(i=0;i (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0];var j=d.createElement(s);var dl=l!='dataLayer'?'&l='+l:'';j.src='//www.googletagmanager.com/gtm.js?id='+i+dl;j.type='text/javascript';j.async=true;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-M677548'); Skip to main content Home About Submit ALERTS / RSS Search for this keyword Advanced Search New Results Impact of Biogenic Structures of the Soil-Nesting Ants Lasius niger and Lasius flavus on the Soil Microarthropod Community in Urban Green Spaces View ORCID Profile Maria Sterzyńska , View ORCID Profile Dariusz J. Gwiazdowicz , View ORCID Profile Paweł Nicia , View ORCID Profile Paweł Zadrożny , View ORCID Profile Gema Trigos-Peral , View ORCID Profile Mohamed W. Negm doi: https://doi.org/10.1101/2025.07.31.667896 Maria Sterzyńska 1 Museum and Institute of Zoology PAS , Twarda 51/55, 00-818 Warsaw, Poland Find this author on Google Scholar Find this author on PubMed Search for this author on this site ORCID record for Maria Sterzyńska For correspondence: majka{at}miiz.waw.pl Dariusz J. Gwiazdowicz 2 Department of Forest Entomology and Pathology, Poznań University of Life Sciences , Wojska Polskiego, 71c, 60-625 Poznań, Poland Find this author on Google Scholar Find this author on PubMed Search for this author on this site ORCID record for Dariusz J. Gwiazdowicz Paweł Nicia 3 Department of Soil Science and Agrophysics, University of Agriculture in Kraków, Al . Mickiewicza 21, 31-120 Kraków, Poland Find this author on Google Scholar Find this author on PubMed Search for this author on this site ORCID record for Paweł Nicia Paweł Zadrożny 3 Department of Soil Science and Agrophysics, University of Agriculture in Kraków, Al . Mickiewicza 21, 31-120 Kraków, Poland Find this author on Google Scholar Find this author on PubMed Search for this author on this site ORCID record for Paweł Zadrożny Gema Trigos-Peral 1 Museum and Institute of Zoology PAS , Twarda 51/55, 00-818 Warsaw, Poland Find this author on Google Scholar Find this author on PubMed Search for this author on this site ORCID record for Gema Trigos-Peral Mohamed W. Negm 4 Department of Plant Protection, Faculty of Agriculture, Assiut University , Assiut 71526, Egypt Find this author on Google Scholar Find this author on PubMed Search for this author on this site ORCID record for Mohamed W. Negm Abstract Full Text Info/History Metrics Preview PDF Abstract Organisms that physically modify their environment, known as ecosystem engineers, can influence resource availability, species interactions and the structure of soil communities. However, the specific effect of ecosystem engineers like ants on the abundance and diversity of non-engineering soil organisms remains understudied. To address this knowledge gap, we conducted a survey of a multi-taxon belowground community of soil microarthropods - Collembola, Mesostigmata, Oribatida and Actinedida - in urban areas, comparing nest mounds of the ant species Lasius niger (Linnaeus, 1758) and Lasius flavus (Fabricius, 1782) with areas without ant-nesting activity (control). We hypothesised differences in abundance and distribution patterns of different soil microarthropod taxa between ant mounds and the control soil. We also hypothesised that ant-induced soil disturbance is species specific, and may result in different patterns of diversity and composition of soil microarthropod assemblages within trophic levels, such as among detritivores (e.g. Collembola) and predators (e.g. Mesostigmata, which are mainly free-living predatory species in soil litter habitats). Our results reveal how ecological filters shape different soil microarthropod groups’ responses to ant-driven changes in their environment. As we expected, soil disturbance caused by ant nest-building activity significantly influenced the abundance, distribution patterns and diversity of these soil microarthropods. 1. Introduction Organisms capable of physically engineering ecosystems have the capacity to directly or indirectly, and either positively or negatively, shape resource availability, species interactions, species diversity and abundance [ 1 ]. Soil engineers, as defined by [ 2 ], can directly or indirectly modify resource availability through bioturbation processes that actively alter the physical state of abiotic and biotic material [ 3 ], and create distinctly specific environmental conditions that differ from those of the surrounding soil [ 4 , 5 ]. Specifically, soil engineers perform continuous physical work on soils and sediments, displacing and transporting soil organic and mineral compounds, and creating functional biogenic structures such as organo-mineral aggregates (e.g. faeces, mounds, aggregates and gallery walls) and macropores (e.g. galleries and chambers). These structures have specific physical, chemical and biological properties [ 6 ], and are therefore considered to be important drivers of soil diversity and ecological filters, potentially influencing the occurrence and distribution of other soil species, including microarthropods. Soil provides habitat for a wide range of organisms varying in size (from micro-to macrofauna), trophic status and ecological function [ 5 , 7 , 8 ]. These organisms contribute to the performance of essential soil-based ecosystem functions and services, such as litter fragmentation and decomposition, nutrient cycling, soil structuring and soil organic matter formation (e.g. [ 5 , 9 ]). Within the soil mesofauna, Collembola and Acari are the two most diverse and abundant groups of soil microarthropods inhabiting the upper soil horizons. As belowground consumers, they share a similar pool of resources and predators, and are members of soil micro-food webs defined by the detritivore and microbivore trophic guilds [ 10 ]. However, little is known about how soil engineers can influence the abundance and diversity of different trophic guilds among soil microarthropods. Biogenic structures and/or microhabitats created by ecosystem engineers are ecologically important because they are considered to be resource and activity hotspots, patches or even islands in soil ecosystems (e.g. [ 4 , 11 ]). As a resource, biogenic structures not only modify the soil structure, but can also influence local community assembly processes that shape the identity and abundance of species in belowground communities. It is widely recognised that the linkages between soil fauna and soil processes and ecosystem functioning depend on biodiversity, and are controlled by both inter- and intraspecific complex interactions, as well as bottom-up and top-down processes [ 12 ]. However, the links between the effects of soil engineers, the biodiversity of non-engineering soil-dwelling species, and ecological interactions in the soil have not been clearly and sufficiently identified. Ants contribute significantly to soil biodiversity and belowground soil processes [ 13 ]. As they can physically engineer the soil ecosystem, they serve an important function as ‘facilitators’ or ‘regulators’ of biotic interactions in the soil [ 14 ]. They also support soil-based ecosystem services [ 15 ] and are one of the main sources of small-scale soil disturbance [ 16 ]. The soil-engineering activity of ants is based on digging to construct nest structures that include chambers in which to host the colony, and galleries and sheets that facilitate foraging activity both below and above the ground. It is generally accepted that the characteristics of the soil and the architecture of the nest structures allow ants to reduce environmental hazards (i.e. predation, and temperature and moisture fluctuation) and optimise colony development [ 4 , 17 , 18 ]. On the other hand, these biogenic structures produced by ant engineering activity—usually referred to as the myrmecosphere [ 19 ]—alter soil physical and chemical properties, nutrient cycling and energy fluxes [ 20 – 24 ]. Consequently, ants alter the ecological filters that shape above- and belowground interactions by creating environments that constrain the species pool by promoting a set of biological traits of organisms that facilitate mutualistic associations with ants, thus supporting species coexistence [ 25 – 27 ]. For example, the engineering activity of ants can increase microbial processes and interactions, which in turn accelerates microbial exchange pools [ 28 – 30 ]. Ant soil engineering can also alter the soil seed bank [, 31 , 32 ]. Nevertheless, there are few studies investigating the effect of different ant species on belowground communities such as soil microarthropods, and most of the existing studies investigated wood ant ( Formica rufa group) nests, which consist of aboveground mounds built with needles, leaves, twigs and bark material [ 33 – 36 ]. However, although some trends may be shared, the diversity and distribution patterns of soil microarthropods in the abovementioned mounds may differ significantly from those of microarthropods inhabiting mounds or small hills of granular soil built by soil-nesting ant species. Despite this, the latter have hardly been studied [ 37 , 38 ]. Consequently, little is known about the impact of soil-nesting ant species on the community structure and diversity patterns of soil microarthropod taxa associated with different trophic groups, such as detritivores and predators. The contribution of ants to improving soil fertility is important for the maintenance of local biodiversity, particularly in highly disturbed habitats such as urban environments, where the soil’s suitability for plant growth may be compromised. In urban ecosystems, anthropogenic disturbances—including polluted precipitation, hydrological drought, changes in resource availability, and conversion to different land uses—result in altered soil conditions (see [3941]). Nevertheless, urban green spaces mostly resemble natural plant communities (e.g. [ 42 ]) and are commonly colonised by ants [ 43 – 45 ]. In this study, we conducted a survey of a multi-taxon belowground community of soil microarthropods—Collembola, Mesostigmata, Oribatida and Actinedida—occurring both in mounds and in the adjacent soil habitats of the two soil-nesting ant species Lasius niger (Linnaeus, 1758) and Lasius flavus (Fabricius, 1782) in urban areas. These two soil-nesting ant species differ greatly in their nesting and foraging habits. L. niger is a semi-predatory species with mostly epigean foraging habits (although it feeds in all biocoenotic layers), and often nests under stones [ 44 ]. L. flavus is a more cryptic species with hypogean foraging habits, feeding mostly on honeydew exuded by root aphids, and its mound nests are often overgrown with mosses, herbs and grasses [ 44 ]. In this study, we hypothesised that due to their different ecology, these two soil-nesting ant species differed in the type of soil disturbance caused by their activity. We further inferred that this would result in differences in the structure and composition of the soil microarthropod communities inhabiting their mounds and the surrounding areas. Our study aimed to identify gaps in the knowledge on biotic filtering in relation to ant–soil–microarthropod interactions, as well as to provide new insights into the influence of ant activity on the soil microarthropod community. Specifically, the main objectives of this study were to assess the effects of soil disturbance by soil-nesting ant species on the density and distribution of higher soil microarthropod taxa, and to evaluate the specific engineering effects of the two soil-nesting ant species L. flavus and L. niger on the diversity patterns and structures of soil microarthropod assemblages with different trophic positions: detritivores and predators, represented by the two taxa Collembola and Mesostigmata, respectively. 2. Materials and Methods 2.1. Study Sites The study was conducted in semi-natural grasslands of open green spaces in the Warsaw agglomeration, where the soil-nesting ants L. niger and L. flavus are the most abundant ant species [ 44 ]. The three sites were located on frequently mowed (5 times a year) green lawns whose vegetation resembled pasture communities (Cynosurion alliance) or wet meadows (Arrhenatherion alliance) belonging to the order Arrhenatheretalia ( Figure 1 ). Each site was geographically referenced using ArcView 9.3. The soils were classified as Cambisols and Luvisols formed from fluvioglacial sands [ 46 ]. Download figure Open in new tab Figure 1. Location of the study area and distribution of the examined semi-natural grassland sites in the urban green space. ( A) — map of the study sites in the open green areas in the Warsaw agglomeration, ( B) – location of the study area in Poland, ( C) – location of the study area in the Warsaw agglomeration. Site 1 – 52° 10’10” N 21° 02’03” E; Site 2 – 52° 10’49” N 21° 02’35” E; Site 3 – 52° 10′12” N 21° 02′58” E. 2.2. Soil Microarthropod Sampling and Processing To cover site variability, samples were taken from six mounds with living ant colonies: three from L. niger mounds and three from L. flavus mounds. A soil sample (5 cm in diameter × 10 cm deep) was taken from the centre of each mound. In addition, a control sample was taken 1 m from each mound in a grassland area with no ant-nesting activity. In total, 72 soil samples from ant mounds (36 from L. niger and 36 from L. flavus ) and 72 soil samples from grasslands (control microhabitat) were collected during the growing season (from June to September 2015). Soil microarthropods from the mounds and adjacent control microhabitats were extracted in a modified MacFadyen high gradient apparatus for a maximum of 10 days. The material was then identified to the higher taxonomic levels of soil microarthropods (Collembola, Mesostigmata, Oribatida and Actinedida). Subsequently, the composition and structure of two assemblages of soil microarthropods, Collembola and Mesostigmata, representing different feeding habits—detritivores and predators, respectively—were assessed for the L. niger and L. flavus mounds. Identification of Collembola species was performed using recent comprehensive monographs [ 47 – 54 ], following the nomenclature of [ 55 ]. The studied material was deposited in the Museum and Institute of Zoology, PAS (Warsaw, Poland). To identify the species of Mesostigmata (Acari), both semi-permanent (using lactic acid) and permanent microslides (using Hoyer’s medium) were prepared. All the mesostigmatic mites were examined with a light microscope (Zeiss Axioskop 2) and identified with reference to taxonomic literature (e.g. [ 56 – 58 ]). 2.3. Abiotic Factors To characterise the microclimatic and soil conditions, we assessed soil moisture (soil volumetric water content, VWC), soil temperature (T soil ) and electrical conductivity (ECe) using automated ProCheck sensors (Decagon Devices, Pullman, USA). Measurements were taken from the central part of all ant mounds and control microhabitats in parallel with soil microarthropod sampling. Soil pH and total carbon, nitrogen, potassium and phosphorus content (hereafter referred to as major elements, quantified in g kg -1 ) were determined in the laboratory from the soil material collected in September 2015. Soil pH was measured using a 1:2.5 soil:water suspension by the potentiometric method. Total carbon (TC) and total nitrogen (TN) concentration were quantified by the total combustion method using a LECO CNS 2000 automatic analyser. The concentrations of total phosphorus (TP) and potassium (TK) were analysed by plasma–optical emission spectrometry after digestion of the samples in a mixture of nitric and perchloric acids [ 59 ], and quantified in g kg -1 . The C/N nutrient ratio was calculated from these data. 2.4. Data Analysis Differences in soil physicochemical properties and the abundance of soil microarthropod taxa (Collembola, Oribatida, Mesostigmata and Actinedida) between soil from mounds of the two soil-nesting ant species L. flavus and L. niger and from control microhabitats (without ant-nesting activity), as well as between L. niger and L. flavus mounds, were compared using a two-sided Mann–Whitney U test. Principal component analysis (PCA) was then used to test whether the ants’ engineering activity associated with changes in soil properties was able to distinguish Lasius spp. mound microhabitats from control soils, as well as showing microhabitat differences between L. niger and L. flavus mounds. To determine the significance of differences between Lasius spp. mound and control microhabitats, and between L. niger and L. flavus mound microhabitats, the PCA results of the first and second axes were used as synthetic dependent variables in the Mann–Whitney U test. The effect of microhabitat type (M; ant mounds and control soil), season (S; spring, autumn) and site location (Sl; site1, site2, site3) on the variation in soil microarthropod taxa (Collembola, Oribatida, Mesostigmata and Actinedida) was assessed by redundancy analysis (total RDA), since the value of gradient length calculated in the detrended correspondence analysis (DCA) for the first axis was shorter than 3.0 standard deviations (SD). Furthermore, the effect of species-specific ant mounds (AM) of L. flavus (LF) and L. niger (LN), season and site location on the variation of Collembola and Mesostigmata assemblages was tested using canonical correspondence analysis (total CCA), as the gradient length exceeded 3 SD (see [ 60 ]). Variation partitioning was performed using partial models, including pRDA-(partial redundancy analysis) and pCCA (partial canonical correspondence analysis)-based models, to explain the unique (conditional) and shared effects of the investigated factors [ 61 ]. The aim of these models was to reflect the relative importance of microhabitat type, season and site location as a group of predictors of variation in soil microarthropod taxa and to represent the combined effect of ant mounds, season and site location on the variation in Collembola and Mesostigmata assemblages’ composition. An interactive forward selection procedure was then used to assess the statistical significance of each individual explanatory variable of the factors studied—microhabitat type (M - ant mound, soil - control), season and site location—to rank the explanatory variables by their importance in determining the significance of their effect on variation in soil microarthropod taxa. The forward selection method was also used in a stepwise regression procedure to predict the significance of each variable for factors such as ant mound type, season and site location on variation in Collembola and Mesostigmata assemblages’ composition. In the model, the adjusted variation was applied using the number of degrees of freedom. A forward selection procedure (interactive) was used with 999 permutations, and the p-value was adjusted by the false discovery rate. The significance of the models was estimated using the unconstrained Monte Carlo permutation test. We performed the multivariate analysis using the data matrix species and/or soil microarthropod taxa × ant mound collected at the site during the same sampling period, and then standardised to square metres. Before the multivariate analyses, data were log(x+1)-transformed to meet assumptions of normality and homogeneity of variance. The significance level for all analyses was set at α = 0.05. Calculations were performed using the software packages Statistica 10.0 and Canoco 5.0. 3. Results 3.1. Soil Properties The descriptive statistics of the recorded soil properties for the Lasius flavus mounds, Lasius niger mounds, the combined ant mounds (mounds of L. niger and L. flavus considered together, hereafter referred to as Lasius spp. and/or LF+LN), and the control microhabitat soils are presented in Table 1 . The two-tailed Mann–Whitney U test showed that the ant mounds (LF+LN) had significantly lower soil moisture (U = 466.5, z = −2.03, N = 36, p = 0.040) and higher total potassium (TK) (U = 421.0, z = 2.77, N = 18, p = 0.006) than the surrounding control grassland soils. However, there were no significant differences in the mean ranks of the other soil characteristics measured between L. flavus (LF) and L. niger (LN) mounds. View this table: View inline View popup Table 1. Summary statistics (mean ± standard deviation) of soil characteristics from the ant mounds of Lasius niger (LN), Lasius flavus (LF), Lasius flavus (LF) + Lasius niger (LN), and control microhabitats (C) without ant-nesting activity in open green space in the Warsaw agglomeration. TK – total potassium content; TP – total phosphorus content; TN – total nitrogen content; TC – total carbon content; EC – electrical conductivity. Values presented in the table are means across replicates of each site for soil moisture, T soil and EC (n = 36), and for TK, TP, TN, TC and C/N ratio (n = 18). The results of the PCA ordination confirmed the differences in soil properties between ant mounds (LF+LN) and control microhabitats (C) without ant-nesting activity, with significantly higher factor loadings for the first PC axis (two-tailed Mann–Whitney U test for sum range of soil properties: p < 0.05; n = 36 in all cases; Figure 2A ). PCA axis 1 accounted for 73.9% of the total variance in soil properties, with variables such as total nitrogen (TN), phosphorus (TP) and potassium (TK) contents having the largest positive loadings; in contrast, soil temperature, followed by soil pH, had the largest negative loadings. PCA axis 2 accounted for 27.0% of the total variance and had the largest positive loading on TK and a negative loading on soil moisture ( Figure 2B , Supplementary Table S1). In the PCA plot ( Figure 2A ), the ant mounds at the top are separated from the control sites at the bottom, but over the season the ant mounds and control sites become less tightly clustered. Download figure Open in new tab Figure 2. The PCA ( A ) ordination biplot (PC 1 and PC 2) and ( B ) correlation biplot (PC 1 and PC 2) of the soil-nesting Lasius flavus + Lasius niger mounds and non-nested (control) plots, with the microhabitat type (ant-mound and control) as the supplementary variables. The model was calculated using raw data of soil characteristics and interspecies correlation scaling, with the species score divided by the standard deviation (SD) and centred by species, not standardised by sample. The results of further PCA ordination points showed differences in soil properties between L. flavus and L. niger mounds, with significantly larger factor loadings for the first PC axis (two-tailed Mann–Whitney U test for sum range of soil properties: p < 0.05; n = 18 in all cases; Figure 3A ). PCA axis 1 accounted for 78.1% of the total variance in soil properties. The highest negative factor loading was found for soil temperature, while the highest positive factor loading was found for TC, TN and TP content. PCA axis 2 accounted for 12.7% of the total variance and had the highest factor loadings positively related to EC and negatively related to TK ( Figure 3B , Supplementary Table S2). Download figure Open in new tab Figure 3. The PCA ( A ) ordination biplot (PC 1 and PC 2) and ( B ) correlation biplot (PC 1 and PC 2) of the soil-nesting Lasius flavus (LF) and Lasius niger (LN) mounds with the microhabitats of ant-mounds as the supplementary variables. The model was calculated using raw data of soil characteristics and interspecies correlation scaling, with the species score divided by the standard deviation (SD) and centred by species, not standardised by sample. Soil-characteristic abbreviations are given in Table 1 . The soils of L. niger mounds were warmer and more fertile (characterised by higher TK, TN, TC and TP contents) than L. flavus mounds. 3.2. Soil Microarthropod Taxonomic Composition and Abundance Patterns Descriptive statistics of the recorded abundances of soil microarthropod taxa from the L. flavus and L. niger mounds and the control (surrounding grassland) soil are presented in Table 2 . A two-tailed Mann–Whitney U test showed that the abundance of soil microarthropods with generalist feeding habits—i.e. Collembola and Oribatida (classified as detritivores)—was significantly reduced within the ant mounds (LF+LN) compared to the control microhabitats (two-tailed Mann–Whitney U test for Collembola: p = 0.0002; Oribatida: p = 0.0009; n = 36 in all cases). Conversely, the abundance of soil microarthropods characterised by specialised feeding habits—i.e. Mesostigmata and Actinedida (classified as predators)—remained constant between the ant mounds and control microhabitats. In contrast, the abundance of soil microarthropod taxa in L. flavus and L. niger mounds did not differ significantly, except for Mesostigmata, which were more abundant in L. niger mounds than in L. flavus mounds (two-tailed Mann–Whitney U test for Mesostigmata: p = 0.007, n = 18) ( Table 2 ). View this table: View inline View popup Table 2. Abundance of soil microarthropod taxa in Lasius flavus (LF), Lasius niger (LN), and Lasius flavus + Lasius niger (LF+LN) ant mounds, and in control microhabitats without ant-nesting activity (C) in open green space in the Warsaw agglomeration. Values are means ± standard deviation (SD) of 18 replicates of taxon abundance in LF and LN ant mounds, and 36 replicates of taxon abundance in LF+LN ant mounds and control microhabitats (C); D – density of soil microarthropod taxa (ind.m -2 ). The results of the total RDA model confirmed that the study factors (the soil microhabitat—i.e. nested ant mound/non-nested control grassland soil—as well as the site and season) accounted for 13.1% of the variation (adjusted explained variation 7.9%). The results of this model explained a significant amount of variation in soil microarthropod components (λ-trace = 0.1306, F-ratio = 2.5, p-value = 0.006). Variation decomposition with the partial RDA model revealed that soil microhabitat (M) and season (S) had the highest unique effects within the model, accounting for 7.1 and 4.4% of the adjusted variation, respectively, while the contribution of the site factor to the total variation was negative (−1.5%) and non-significant. Furthermore, the joint overlap of all three factors was relatively small (p < 0.1%; Table 3 ). View this table: View inline View popup Table 3. Abundance of soil microarthropod taxa in Lasius flavus (LF), Lasius niger (LN), and Lasius flavus + Lasius niger (LF+LN) ant mounds, and in control microhabitats without ant-nesting activity (C) in open green space in the Warsaw agglomeration. Values are means ± standard deviation (SD) of 18 replicates of taxon abundance in LF and LN ant mounds, and 36 replicates of taxon abundance in LF+LN ant mounds and control microhabitats (C); D – density of soil microarthropod taxa (ind.m -2 ). Further analysis using partial model RDA with interactive forward selection identified the significant effects of soil-nesting ant mound, control site and season (spring and autumn) as single variables on the variation of soil microarthropod communities ( Table 4 ). View this table: View inline View popup Download powerpoint Table 4. Results of interactive forward selection in RDA and ranking of examined factors on variation in the soil microarthropod communities. Model calculated with log(x+1) transformed data; pseudo-F – F statistic; p-value – significance level of the effect tested by Monte Carlo permutation test with 999 permutations; p-value (adj.) – correction with false discovery rate. This analysis also showed that the presence of soil-nesting mounds of both ant species had a negative influence on the distribution pattern of soil microarthropod taxa ( Figure 4 ). The results of subsequent RDA and partial RDA analyses showed that the abundance of soil microarthropods did not respond to environmental changes (soil disturbance) in L. niger and L. flavus ant mounds (Table S3). Download figure Open in new tab Figure 4. Distribution pattern of soil microarthropod taxa within soil-nesting ant mounds and non-nested (control) sites. RDA model calculated with log(x+1) transformed data. Coll – Collembola; Orib – Oribatida; Mes – Mesostigmata; Acti – Actinedida. 3.3. Diversity Pattern and Structure of Collembola Assemblages A total of 25 Collembola species were recorded in the ant mounds of L. niger and L. flavus (Table S4). The majority of species recorded were soil-dwelling, while the myrmecophilous species were represented by only two species, Cyphoderus albinus and C. bidenticulatus . The effects of L. flavus and L. niger ant mounds on the mean ranks of collembolan abundance (A), species richness (S) and diversity index (H’) were not statistically significant (two-tailed Mann–Whitney U test (N = 18, p > 0.05 in all cases). The total CCA analysis showed that all the investigated factors—LN vs LF ant mounds, season and site—accounted for 17.4% of the variation (adjusted explained variation 3.6%) in the species dataset (λ-trace = 1.98, F-ratio = 1.30, p-value = 0.02). Variation decomposition ( Table 5 ) showed that season had the greatest unique effect, accounting for 3.1% of the adjusted variation, while ant mounds and site were not significant and explained less of the variation (0.7% and 0.6%, respectively). The results of the partial CCA with site as a covariate showed further significant differences in Collembola assemblage composition between ant mounds (λ-trace = 0.702, F-ratio = 1.481, p-value = 0.008). For example, during the autumn season, myrmecophilous collembolan species such as C. bidenticulatus and C. albinus were observed to be associated with L. flavus mounds (Figure S1). View this table: View inline View popup Table 5. Variance partitioning among ant mounds (AM: Lasius niger and Lasius flavus ), site (Sl) and season (S) on variation in Collembola assemblages from soils in green spaces in Warsaw. Conditional effect performed by CCA and partial CCA model. DF – degree of freedom; mean square – denominator of F-statistic; pseudo-F – F statistic; p-value – significance level of the effect tested by Monte Carlo permutation test; CCA model calculated with log(x+1) transformed data. 3.4. Diversity Pattern and Structure of Mesostigmata Assemblages A total of 21 Mesostigmata species were recorded in the ant mounds of L. niger and L. flavus (Table S5). Most of the species recorded were soil-dwelling. The effect of L. flavus and L. niger ant mounds on the mean ranks of Mesostigmata abundance (A) was significant ( Table 2 ), whereas the species richness (S) and diversity index (H’) were not statistically significant (two-tailed Mann–Whitney U test (N = 18, p > 0.05 in all cases). The total CCA analysis showed that all predictors included in the model accounted for 15.9% of the variation (adjusted explained variation 0.6%), but the model was not significant (λ-trace = 1.475, F-ratio = 1.0, p-value = 0.372). Further variation decomposition and partial CCA analysis with site as a covariate were not significant ( Table 6 , Figure S2). View this table: View inline View popup Table 6. Variance partitioning among ant mounds (AM), site (Sl) and season (S) on variation in Mesostigmata assemblages from soils in green space in Warsaw. Conditional effect performed by CCA and partial CCA model. DF – degree of freedom; mean square – denominator of F-statistic; pseudo-F – F statistic; p-value – significance level of the effect tested by Monte Carlo permutation test; CCA model calculated with log(x+1) transformed data. 4. Discussion One of the main objectives of the present study was to assess the impact of soil-nesting ants’ biogenic structures on belowground communities, with a particular focus on the soil microarthropod taxa Collembola, Oribatida, Mesostigmata and Actinedida. To achieve this goal, the study covered three semi-natural grasslands in the Warsaw metropolitan area, where the abundance and distribution of soil microarthropods were investigated in the mounds of two different species of soil-nesting ants— Lasius flavus (LF) and Lasius niger (LN)—as well as in grassland soils with no ant-nesting activity (control). We also investigated the species-specific effects of the biogenic structures of these two ant species on the diversity and structure of soil microarthropod assemblages at different trophic levels of the soil food web: decomposers (Collembola) and predators (Mesostigmata). As expected, our results showed that ants can shape soil microarthropod assemblages. These results are consistent with other studies showing that ant-induced soil disturbance can cause changes in belowground communities [ 16 , 25 , 33 , 36 ]. The effect of ant ecological pressure on the community structure of soil microarthropods in both types of microhabitats studied (soil-nesting ant mounds vs control soil) was confirmed by the differences found in the abundance of soil microarthropods at the coarse taxonomic level ( Table 2 ). However, our study showed that this effect varied according to the position of the soil microarthropod taxon in the soil food web. In particular, ant pressure had a negative effect on the abundance of taxa with decomposition/detritivore functions in soil (i.e. Collembola and Oribatida) and a neutral effect on the abundance of taxa considered to be top predators in the soil mesofauna food web, i.e. Mesostigmata [ 10 ] ( Table 2 ). This result confirms the importance of edaphic microarthropod food specialisation as a key factor influencing community structure and function [ 62 ], even in microhabitats with ant-induced soil disturbance. Furthermore, we found significant variability in soil microarthropod communities between the studied microhabitats— with and without ant-nesting activity—using the RDA and pRDA models ( Tables 3 , 4 , Figure 4 ), as well as the significant effect of seasonal variation on these communities. These results suggest that the selective filter shaped by ants through which soil microarthropod taxa pass—in addition to exposure to chronic predation—is related not only to disturbance of the soil environment within mounds as a result of ant-nesting activity, but also to variation in microhabitat conditions between mounds and control soils over time. Several authors have indicated that ants, through their nesting activity, directly and indirectly alter both biotic and abiotic properties of the soil environment within the mound [ 1 , 20 , 21 , 63 , 64 ], including soil physicochemical properties or resource availability [ 24 , 25 , 63 – 65 ]. Ant-digging activity also induces structural disturbances associated with fine-scale landscape-level changes, which enhance basic physical processes such as water infiltration and nutrient redistribution [ 66 ]. Our results showed that the negative response of soil microarthropod communities to soil disturbance by soil-nesting ants was mainly associated with significantly increased soil dryness and significantly higher contents of soil-fertility-enhancing macronutrients such as potassium and phosphorus in mound soils compared to control soils without ant-nesting activity ( Table 1 , Table S1, Figure 2 a,b ). Soil microarthropods are highly sensitive to environmental changes caused by drought and soil warming [ 67 – 72 ]. Disturbances in soil physicochemical properties, such as soil drought/temperature and/or soil fertility, can strongly influence their community structure through bottom-up effects [ 70 , 72 – 74 ]. In addition, changes in hydration status, which is a key descriptor of soil physical structure and an important driver of trophic interactions in belowground communities [ 74 ], can locally facilitate predator–prey interactions, leading to a sharp decrease in the abundance of prey communities. Conversely, changes in microhabitat conditions in the nests of L. niger and L. flavus ants due to their activity, particularly those associated with increases in soil fertility, did not affect the abundance of Mesostigmata ( Table 2 ). Mesostigmata is a taxon considered to be an important predator in the food web of soil mesofauna [ 75 ]. This result contradicts the study by [ 73 ], who found an increased proportion of predatory soil microarthropods in temperate grassland communities with improvements in soil fertility. The present results are also inconsistent with the study of [ 36 ], who reported a negative effect of red ants ( Formica polyctena ) on microarthropod taxa in a semi-arid pine forest. These discrepancies may be due to different degrees of soil pH disturbances influenced by the specific ant species, which in turn affects soil-associated biodiversity. Red ants have been shown to alter soil pH, shifting acidic and neutral soils to more alkaline soils, and strongly alkaline soils to more acidic soils [ 77 ]. However, our study found no significant differences in soil pH between areas with Lasius spp. and areas without ant-nesting activity. Furthermore, the present study differs from that of [ 36 ] in terms of the composition of the organisms studied, as the present study included soil consumers of the same size class and within the same microhabitat network [ 10 , 75 ]. Despite the absence of differences in the abundance of Mesostigmata mites in the Lasius spp. vs control microhabitats studied, these organisms may still exert a cascading effect on mesofaunal detritivore taxa. This finding suggests that the density of soil microarthropods in ant nests is not predominantly controlled by bottom-up forces, but rather by top-down regulation. Nevertheless, it is noteworthy that the nest mounds of L. niger and L. flavus did not show species-specific differences in the abundance and occurrence of higher soil microarthropod taxa, except for Mesostigmata, which occurred at much higher densities in the mounds of L. niger ( Table 2 ). This scenario may be related to the different impact of the two species on soil properties due to their different ecological characteristics: L. niger mounds were warmer, more fertile and slightly more humid than L. flavus mounds ( Table 1 . Table S2 Figure 3b ). The differences in Mesostigmata abundance between L. flavus and L. niger mounds revealed in our study are in good agreement with published results showing that moisture and temperature within ant mounds are important for soil microarthropod communities [ 35 ]. 4.1. Effect of L. flavus and L. niger on Collembola Assemblage Composition, Abundance and Distribution Patterns The present study showed that microhabitat variation between L. flavus and L. niger mounds did not significantly affect differences in Collembola abundance, species richness or Shannon diversity index values. However, a more detailed analysis of the Collembola assemblages inhabiting these ant nests suggests that seasonal variation may play an important role in shaping their assemblages. Analysis of the species composition of the Collembola assemblages found in the nests of the two ant species L. flavus and L. niger indicates that most of the assemblages are species that prefer (inhabit) the litter and soil layers. A similar pattern was observed in the Collembola assemblages found in the anthills of Formica s. str. nests [ 34 ]. However, it is noteworthy that the anthills of L. flavus and L. niger also host two additional myrmecophilous species, Cyphoderus albinus and C. bidenticulatus . Despite the well-documented association of the genus Cyphoderus (Nicolet, 1842) with the nests of social insects—including termites, bees and ants (see [ 55 , 78 ])— C. bidenticulatus was found only in L. flavus nesting mounds (Table S4, Figure S1), suggesting that the microhabitat conditions created by this ant species may be particularly conducive to its presence. The variance partitioning results also showed that the variation in Collembola assemblages in L. niger and L. flavus mounds was significantly influenced by seasonal changes (see Table 5 ). The present study did not assess the extent to which seasonal changes in habitat conditions occur in the mounds of both ant species. However, it is recognised that seasonal changes in grassland ecosystems can induce a response in soil microarthropod assemblages, including those of Collembola, and that these changes are mainly related to variations in temperature and precipitation [ 70 , 71 ]. Nevertheless, it is noteworthy that despite the lack of significant variability in Collembola assemblages associated with the unique microhabitats of L. niger and L. flavus , the drier and warmer microhabitats found in L. niger nest mounds (compared to L. flavus mounds) favoured the occurrence of xerophilic Collembola species such as Isotomodes productus and Hemisotoma thermophila . We also found higher concentrations of key soil-fertility components (C, N, K and P) in L. niger mounds, creating a microhabitat favourable for the presence of species associated with habitats rich in decomposing organic matter, such as Proisotoma minima (see [ 51 ]). However, the distribution pattern of Collembola species observed in L. flavus and L. niger mounds may be impacted by variables outside the scope of this study. For example, it is important to note that the species identified in this study have been selected for by urban environmental conditions, and that the colonisation of soil-nesting ant mounds by Collembola assemblages may be affected by both anthropogenic and natural disturbances. 4.2. Effect of L. flavus and L. niger on Mesostigmata Assemblage Composition, Abundance and Distribution Patterns The present study showed that microhabitat variation between L. flavus and L. niger mounds had a significant influence on the abundance of Mesostigmata mite assemblages, but it did not show a discernible effect on changes in mite diversity (see Table S5 and Figure S2). However, it is noteworthy that the studied ant mounds contained almost half as many Mesostigmata mite species as the 53 and 44 species previously identified in L. flavus and L. niger nests in Poland [ 37 , 79 ]. A review of over 40 years of research in Poland on Mesostigmata mites in L. flavus and L. niger nests [ 80 – 82 ] shows high species diversity, with each study typically including a different list of species. This diversity is probably due to the function of the soil as a microhabitat where nests are built, often in humus or mineral layers, supporting numerous mite species. Consequently, the lower diversity of Mesostigmata mites observed in our study may be due to the effects of urban disturbance on these communities (e.g. [ 83 ]). Changes in the microhabitat characteristics of the ant mounds of the studied Lasius species are also reflected in the presence of Mesostigmata species frequently found in the mineral soil layer (e.g. Rhodacarellus silesiacus, Rhodacarus mandibularis ) and in the upper litter layer (e.g. Gamasellodes bicolor, Veigaia nemorensis ). In contrast to the myrmecophilous Mesostigmata found in the nest mounds of red-headed ants ( Formica rufa and F. polyctena ) [84], only Cosmolaelaps vacua and Oplitis conspicua —two soil-dwelling Mesostigmata species often associated with ant nests—were found in the nest mounds of L. niger and L. flavus [ 79 ]. 5. Conclusions Our research has identified the role of ecological filters in the response of soil microarthropod taxa to ant engineering activity. As we expected, soil disturbance caused by ant engineering activity can influence the response of soil microarthropod taxa, including their abundance and distribution patterns. Digging and mound building by the two ant species studied ( L. niger and L. flavus ) not only increased the local heterogeneity of grassland communities in urban green spaces, but also induced soil disturbance that affected the variability of soil microarthropod assemblages. However, species-specific disturbance and the associated change in microhabitat conditions in the mounds of the two ant species studied did not significantly affect the variation in diversity patterns and species composition of the Collembola and Mesostigmata assemblages. In contrast, the results of the present study suggest that seasonal variation in resource availability in mounds and interspecific interactions may be important factors, especially in the assembly of detritivorous—but not predatory—guilds of soil microarthropods. However, it should be noted that while urban green spaces tend to resemble natural plant communities, anthropogenic disturbances (e.g. pollutant deposition, hydrological droughts, changes in resource availability and conversion to different land uses) also strongly modify soil [see 41], including that of the myrmecosphere. Consequently, the response exhibited by the local soil microarthropod community to ant engineering activity in urban areas may be either universal or unique. Footnotes MS e-mail: majka{at}miiz.waw.pl , GT-P e-mail: gtrigs{at}miiz.waw.pl e-mail: dariusz.gwiazdowicz{at}up.poznan.pl e-mail: rrnicia{at}cyf-kr.edu.pl , pawel.zadrozny{at}urk.edu.pl e-mail: waleednegm{at}yahoo.com References 1. ↵ Jones , C.G. ; Lawton , J.H. ; Shachak , M. Positive and negative effects of organisms as physical ecosystem engineers . Ecology 1997 , 78 , 1946 – 1957 . OpenUrl CrossRef Web of Science 2. ↵ Lavelle , P. Faunal activities and soil processes: adaptive strategies that determine ecosystem function . Advances in Ecological Research 1997 , 21 , 93 – 132 . OpenUrl 3. ↵ Taylor , A.R. ; L. Lenoir L .; Vegerfors , B. ; Persson T. Ant and earthworm bioturbation in cold-temperate ecosystems . Ecosystems 2019 , 22 , 981 – 994 . OpenUrl 4. ↵ Jouquet , P. , Dauber , J. , Lagerlöf , J. , Lavelle , P. , Lepage , M. Soil invertebrates as ecosystem engineers: intended and accidental effects on soil and feedback loops . Applied Soil Ecology 2006 , 32 , 153 – 164 . OpenUrl 5. ↵ Paul , E.A. Coleman , D.C. ; Wall , D.H. Soil Fauna: Occurrence, biodiversity and role in ecosystem function . In Soil Microbiology, Ecology and Biochemistry , 4th ed.; Paul , E.A. , Ed.; Elsevier : Amsterdam, The Netherlands 2024 ; pp. 111 – 149 . 6. ↵ Culliney T. Role of Arthropods in maintaining soil fertility . Agriculture 2013 , 3 , 629 – 659 . OpenUrl 7. ↵ Lameed , G.A. Menta , C. Soil fauna diversity – function, soil degradation, biological indices, soil restoration . In Biodiversity Conservation and Utilization in a Diverse World ; Lameed , G.A. , Ed.; IntechOpen : Rijeka, Croatia , 2012 . 8. ↵ Briones , M.J. Soil fauna and soil functions: A jigsaw puzzle . Front. Environ. Sci . 2014 , 2 , 7 . doi: 10.3389/fenvs.2014.00007 OpenUrl CrossRef 9. ↵ Lavelle , P. ; Decäens , T. ; Aubert . M. ; Barot , S. ; Blouin , M. ; Bureau , F. ; Margerie , P. ; Mora , P. ; Rossi , J-P. Soil invertebrates and ecosystem services . European Journal of Soil Biology 2006 , 42 , 3 – 15 . OpenUrl 10. ↵ Potapov , M. ; Beaulieu , F. ; Birkhofer , K. ; Bluhm , S.L. ; Degtyarev , M.I. ; Devetter , M. ; Goncharov , A.A. ; Gongalsky , K.B. ; Klarner , B. ; Korobushkin , D.I. ; Liebke , D.F. ; Maraun , M. ; Mc Donnell , R.J.; Pollierer , M. ; Schefer , I. ; Shrubovych , J. ; Semenyuk , I.J. ; Sendra , A. ; Turna , J. ; Tumova , M. ; Vassilieva , A.B. ;, Chen , T-W. ; Geisen , S. ; Schmidt , O. ; Tiunov , A.V. ; Scheu S. Feeding habits and multifunctional classification of soil associated consumers from protists to vertebrates . Biol. Rev . 2022 , 97 , 1057 – 1117 . OpenUrl CrossRef 11. ↵ Klimek , B. ; Poliwka-Modliborek , H. ; Grześ , I.M. Ant nests as a microbial hot spots in a long-term heavy metal-contaminated soils . Environ Sci Pollut Res 2022 , 29 : 10848 – 10857 . doi: 10.1007/s11356-021-16384-y OpenUrl CrossRef 12. ↵ Bardgett , R. , van der Putten , W. Belowground biodiversity and ecosystem functioning . Nature 2014 , 515 , 505 – 511 . doi: 10.1038/nature13855 OpenUrl CrossRef PubMed 13. ↵ Wills , B. D. ; Landis , D. A. The role of ants in north temperate grasslands: A review . Oecologia 2018 , 186 : 323 – 338 . OpenUrl CrossRef PubMed 14. ↵ Hillel , D. Winsome , T. Fauna . In Encyclopedia of Soils in the Environment ; Hillel , D. , Ed.; Elsevier : Oxford, UK , 2005 ; Volume 1 , pp. 539 – 548 . OpenUrl 15. ↵ Jónsson , J. Ö G. ; Davidsdóttir , B. Classification and valuation of soil ecosystem services . Agricultural Systems 2016 , 145 , 866 – 877 . OpenUrl 16. ↵ Farji-Brener , A.G. ; Werenkraut , V. The effects of ant nests on soil fertility and plant performance: a meta-analysis . J. Anim. Ecol . 2017 , 86 : 866 – 877 . OpenUrl PubMed 17. ↵ Turner , J.S. The Soul of the Superorganism . In The Extended Organism: The Physiology of Animal-Built Structures ; Harvard University Press: Cambridge, MA, USA; London, UK , 2000 ; pp. 179 – 200 . 18. ↵ Odling-Smee , F.J. ; Laland , K.N. ; Feldman , M.W. Niche Construction: The Neglected Process in Evolution ; Princeton University Press : Princeton, NJ, USA , 2003 . 19. ↵ Lavelle , P. Functional domains in soils . Ecological Research 2002 , 17 : 441 – 450 . OpenUrl 20. ↵ Holec , M. ; Frouz , J. The effect of two ant species Lasius niger and Lasius flavus on soil properties in two contrasting habitats . European Journal of Soil Biology 2006 , 42 , 213 – 217 . OpenUrl 21. ↵ Frouz , J. ; Jilková , V. The effect of ants on soil properties and processes (Hymenoptera: Formicidae) . Myrmecological News 2008 , 11 , 191 – 199 . OpenUrl 22. Cammeraat , E.L H. ; Risch A.C. The impact of ants on mineral soil properties and processes at different spatial scales . Journal of Applied Entomology 2008 , 132 , 285 – 294 . OpenUrl 23. Haitao , W. ; Donghui , W. ; Xianguo , L. ; Xiaomin , Y. Spatial distribution of ant mounds and effects on soil physical properties in wetlands of the Sanjiang plain, China . Acta Ecologica Sinica 2010 , 30 , 270 – 275 . OpenUrl 24. ↵ Bierbaß , P. ; Gutknecht , J.L.M. ; Michalzik , B. Nest-mounds of the yellow meadow ant (Lasius flavus) at the “Alter Gleisberg”, Central Germany: Hot or cold spots in nutrient cycling? Soil Biol. Biochem . 2015 , 80 , 209 – 217 . doi: 10.1016/j.soilbio.2014.09.020 . OpenUrl CrossRef 25. ↵ De Almeida , T. ; Mesiéard , F. ; Santonja , M. ; Gros , R. ; Dutoit , T. ; Blight , O. Above- and below-ground effects of an ecosystem engineer ant in Mediterranean dry grasslands . Proc. R. Soc. B . 2020 , 287 , 20201840 . OpenUrl PubMed 26. Folgarait , P.F. Ant biodiversity and its relationship to ecosystem functioning: a review . Biodiversity and Conservation 1998 , 7 , 1221 – 1244 . OpenUrl 27. ↵ Capinera , J.L. Danoff-Burg , J.A. Myrmecophiles . In Encyclopedia of Entomology ; Capinera , J.L. , Ed.; Springer : Dordrecht, The Netherlands , 2008 . doi: 10.1007/978-1-4020-6359-6_4764 OpenUrl CrossRef 28. ↵ Dauber , J. ; Wolters , V. Microbial activity and functional diversity in the mounds of three different ant species . Soil Biology and Biochemistry 2000 , 32 , 93 – 99 . OpenUrl 29. Dauber , J. ; Niechoj , R. ; Baltruschat , H. ; Wolters , V. Soil engineering ants increase grass root arbuscular mycorrhizal colonization . Biology and Fertility of Soils 2008 , 44 , 791 – 796 OpenUrl 30. ↵ Boots , B. ; Clipson , N. Linking ecosystem modification by the yellow meadow ant (Lasius flavus) to microbial assemblages in different soil environments . European Journal of Soil Biology 2013 , 55 , 100 – 106 . OpenUrl 31. ↵ Dauber , J. ; Rommeler , A. ; Wolters , V. The ant Lasius flavus alters the viable seed bank in pastures . European Journal of Soil Biology 2006 , 42 , 157 – 163 . OpenUrl 32. ↵ O’Grady , A. ; Breen , J. ; Harrington , T.J. ; Courtney , R. The seed bank in soil from the nests of grassland ants in a unique limestone grassland community in Ireland . Ecological Engineering 2013 , 61 , 58 – 64 . OpenUrl 33. ↵ Laakso , J. ; Setälä , H. Composition and trophic structure of detrital food web in ant nest mounds of Formica aquilonia and in the surrounding forest soil . Oikos 1998 , 81 , 266 – 278 . OpenUrl 34. ↵ Sleptzova , E.V. ; Reznikova , Z.I. Formation of springtail (Collembola) communities during colonization of ant-hills . Entomological Review 2006 , 86 : 373 – 382 . OpenUrl 35. ↵ Elo , R.A. ; Penttinen , R. ; Sorvari , J. A comparative study of oribatid mite communities in red wood ant Formica polyctena nests and surrounding soil in a Finnish oak forest . Insect Conservation and Diversity 2016 , 9 ( 3 ), 210 – 223 . doi: 10.1111/icad.12159 . OpenUrl CrossRef 36. ↵ Çakur , M. ; Akburak , S. ; Sargıncı , M. The negative effect of wood ants (Formica rufa) on microarthropod density and soil biological quality in a semi-arid pine forest . Pedobiologia 2019 , 77 , 150593 . doi: 10.1016/j.pedobi.2019.150593 . OpenUrl CrossRef 37. ↵ Kamczyc , J. ; Gwiazdowicz , D.J. The diversity of soil mites (Acari: Mesostigmata) in yellow ant (Lasius flavus) nests along a gradient of land use . Biologia 2013 , 68 ( 2 ), 314 – 318 . doi: 10.2478/s11756-013-0154-x OpenUrl CrossRef 38. ↵ Arroyo , J. ; O’Grady , A. ; Vance , H. ; Bolger , T. The mite (Acari: Oribatida, Mesostigmata) assemblages associated with Lasius flavus (Hymenoptera: Formicidae) nests and surrounding soil in an Irish grassland . Biology and Environment: Proceedings of the Royal Irish Academy 2015 , 115B , 1 – 12 . OpenUrl 39. Scharenbroch , B.C. ; Lloyd , J.E. ; Johnson-Maynard , J.L. Distinguishing urban soils with physical, chemical, and biological properties . Pedobiologia 2005 , 49 ( 4 ), 283 – 296 . doi: 10.1016/j.pedobi.2004.12.002 OpenUrl CrossRef Web of Science 40. Pickett , S.T.A. ; Cardenasso , M.L. Altered resources, disturbance, and heterogeneity: A framework for comparing urban and non-urban soils . Urban Ecosyst . 2009 , 12 , 23 – 44 . doi: 10.1007/s11252-008-0047-x OpenUrl CrossRef 41. Pickett , S.T.A. ; Cadenasso , M.L. ; Grove , J.M. ; Boone , C.G. ; Groffman , P.M. ; Irwin , E. ; Kaushal , S.S. ; Marshall , V. ; McGrath , B.P. ; Nilon , C.H. ; Pouyat , R.V. ; Szlavecz , K. ; Troy , A. ; Warren , P. Urban ecological systems: Scientific foundations and a decade of progress . Journal of Environmental Management 2011 , 92 ( 3 ), 331 – 362 . doi: 10.1016/j.jenvman.2010.08.022 OpenUrl CrossRef PubMed Web of Science 42. ↵ Kelcey J.G. , Müller N. Sudnik-Wójcikowska , B. , Galera H. Warsaw . In Plants and habitats of European cities . Kelcey J.G. , Müller N. , Eds.; Springer , New York , 2011 , pp 499 – 545 . 43. ↵ Ślipiński , P. ; Żmihorski , M. ; Czechowski , W. Species diversity and nestedness of ant assemblages in an urban environment . Eur. J. Entomol . 2012 , 109 , 197 – 206 . doi: 10.14411/eje.2012.026 OpenUrl CrossRef 44. ↵ Czechowski , W. ; Radchenko , A. ; Czechowska , W. ; Vepsäläinen , K. The Ants (Hymenoptera: Formicidae) of Poland with Reference to the Myrmecofauna of Europe (Fauna Poloniae n.s.4 ). Natura Optima Dux Foundation :: Warsaw , Poland , 2012 . 45. ↵ Trigos-Peral , G. ; Rutkowski , T. ; Witek , M. ; Ślipiński , P. ; Babik , H. ; Czechowski , W. Three categories of urban green areas and the effect of their different management on the communities of ants, spiders and harvestmen . Urban ecosystems , 2020 , 23 , 803 – 818 . doi: 10.1007/s11252-020-00949-9 OpenUrl CrossRef 46. ↵ IUSS Working Group WRB . World Reference Base for soil resources a framework for international classification and communication, World Soil Resource Reports , Vol. 103 . FAO, ISRIOC, IUSS , Rome , 2006 ; pp. 1 – 116 . OpenUrl 47. ↵ Fjellberg , A. The Collembola of Fennoscandia and Denmark . Part I: Poduromorpha; Fauna Entomologica Scandinavica , Vol. 35 ; Brill : Leiden, The Netherlands , 1998 ; pp. 1 – 183 . OpenUrl 48. Fjellberg , A. The Collembola of Fennoscandia and Denmark . Part II: Entomobryomorpha and Symphypleona; Fauna Entomologica Scandinavica , Vol. 42 ; Brill : Leiden, The Netherlands , 2007 ; pp. 1 – 264 . OpenUrl 49. Pomorski , R.J. Onychiurinae of Poland (Collembola: Onychiuridae); Genus, Supplement 9; Museum and Institute of Zoology PAS : Warsaw, Poland , 1998 ; pp. 1 – 201 . 50. Bretfeld , G. Symphypleona . Synopses on Palaearctic Collembola; Abhandlungen und Berichte des Naturkundemuseums Görlitz , 1999 , 71 , 1 – 318 . OpenUrl 51. ↵ Potapov , M. Isotomidae . Synopses on Palaearctic Collembola; Abhandlungen und Berichte des Naturkundemuseums Görlitz , 2001 , 73 , 1 – 603 . OpenUrl 52. Dunger , W. Thibaud , J.-M. ; Schulz , H.-J. ; da Gama Assalino , M.M. Hypogastruridae. In Synopses on Palaearctic Collembola , Vol. 4 : Hypogastruridae ; Dunger , W. , Ed.; State Museum of the Natural History Museum of Görlitz : Görlitz , Germany , 2004 ; pp. 1 – 287 . OpenUrl 53. Dunger , W. ; Schlitt , B. Tulbergiidae . Synopses on Palaearctic Collembola; Soil Organisms , 2011 , 83 , 1 – 168 . OpenUrl 54. ↵ Jordana , R. Capbryinae and Entomobryini . Synopses on Palaearctic Collembola; Soil Organisms , 2012 , 84 , 1 – 390 . OpenUrl 55. ↵ Bellinger , P.F. ; Christiansen , K.A. ; Janssens , F. Checklist of the Collembola of the World ; 1996 – 2020 . Available online: http://www.collembola.org/ (accessed on 29 May 2020 ). 56. ↵ Karg , W. Acari (Acarina), Miben Parasitiformes (Anactinochaeta ), Cohors Gamasina Leach; Gustav Fischer Verlag : Jena, Germany , 1993 . 57. Mašán , P. Mites of the Cohort Uropodina (Acarina, Mesostigmata) in Slovakia; Annotationes Zoologicae et Botanicae , 2001 , 223 , 1 – 320 . OpenUrl 58. ↵ Gwiazdowicz , D.J. Ascid Mites (Acari, Mesostigmata) from Selected Forest Ecosystems and Microhabitats in Poland; Wydawnictwo Akademii Rolniczej : Poznań, Poland , 2007 . 59. ↵ Ostrowska , A. ; Gawliński , S. ; Szczubiałka , Z. Metody analizy i oceny właściwości gleb i roślin . Katalog ; Instytut Ochrony Środowiska : Warszawa, Poland , 1991 . (In Polish) 60. ↵ Lepš , J. ; Šmilauer , P. Multivariate Analysis of Ecological Data Using CANOCO ; Cambridge University Press : Cambridge, UK , 2003 . 61. ↵ Borcard , D. ; Legendre , P. ; Drapeau , P. Partialling out the spatial component of ecological variation . Ecology 1992 , 73 , 1045 – 1055 . OpenUrl CrossRef Web of Science 62. ↵ Velazco , V.N. ; Saravia , L.A. ; Coviella , C.E. ; Falco , L.B. Trophic resources of the edaphic microarthropods: A worldwide review of the empirical evidence . Heliyon 2023 , 9 ( 10 ), e20439 . OpenUrl 63. ↵ Sankovitz , M. ; Purcell , J. Ant nests differentially affect soil chemistry across elevational gradients . Insectes Sociaux 2022 , 69 , 293 – 298 . doi: 10.1007/s00040-022-00869-1 . OpenUrl CrossRef 64. ↵ Zhou , W. ; Zhan , P. ; Zeng , M. ; Chen , T. ; Ahang , X. ; Yang , G. ; Guo , Y. Effects of ant bioturbation and foraging activities on soil mechanical properties and stability . Global Ecology and Conservation 2023 , 46 , e02575 . doi: 10.1016/j.gecco.2023.e02575 OpenUrl CrossRef 65. ↵ Ehrle , A. ; Kolle , O. ; Tischer , A. ; Trumbore , S.E. ; Michalzik , B. Effects of mound building Lasius flavus on organic carbon and nutrient fluxes in soils of temperate grassland ecosystems . Pedobiologia 2021 , 84 , 150701 . doi: 10.1016/j.pedobi.2020.150701 OpenUrl CrossRef 66. ↵ Nkem , J.N. ; Lobry de Bruyn , L.A. ; Grant , C.D. ; Hulugalle , N.R. The impact of ant bioturbation and foraging activities on surrounding soil properties . Pedobiologia 2000 , 44 ( 5 ), 609 – 621 . doi: 10.1078/S0031-4056(04)70075-X . OpenUrl CrossRef Web of Science 67. ↵ Kardol P. , Reynolds W.N. , Norby R.J. , Classon A.T. Climate change effects on soil microarthropod abundance and community structure . Applied Soil Ecology 2011 , 47 : 37 – 44 OpenUrl 68. Wu , T. ; Su , F. ; Han , H. ; Du , Y. ; Yu , C. ; Wan , S. Responses of soil microarthropods to warming and increased precipitation in a semiarid temperate steppe . Appl. Soil Ecol . 2014 , 84 , 200 – 207 . doi: 10.1016/j.apsoil.2014.07.0031 OpenUrl CrossRef 69. Liu , J.-L. ; Li , F.-R. ; Liu , L.-L. ; Yang , K. Responses of different Collembola and mite taxa to experimental rain pulses in an arid ecosystem . Catena 2017 , 155 , 53 – 61 . OpenUrl 70. ↵ Xu , G.-L. ; Kuster , T.M. ; Günthardt-Goerg , M.S. ; Dobbertin , M. ; Li , M.-H. Seasonal exposure to drought and air warming affects soil Collembola and mites . PLoS ONE 2012 , 7 ( 8 ), e43102 . doi: 10.1371/journal.pone.0043102 OpenUrl CrossRef PubMed 71. ↵ Guo , Z. ; Chang , H. ; Liu , R. Response of soil microarthropod community to seasonal changes in Urat Desert Steppe, Inner Mongolia . Front. Environ. Sci . 2022 , 10 , 893913 . doi: 10.3389/fenvs.2022.893913 . OpenUrl CrossRef 72. ↵ Flórián , N. , Ladányi , M. ; Ittzés , A. ; Kröel-Dulay , G. ; Ónodi , G. ; Mucsi , M. ; Szili-Kovács , T. ; Gergócs , V. ; Dányi , L. ; Dombos , M. Effects of single and repeated drought on soil microarthropods in a semi-arid ecosystem depend more on timing and duration than drought severity . PLoS ONE 2019 , 14 ( 7 ), e0219975 . doi: 10.1371/journal.pone.0219975 OpenUrl CrossRef PubMed 73. ↵ Cole , L. ; Buckland , S.M. ; Bardgett , R.D. Relating microarthropod community structure and diversity to soil fertility manipulations in temperate grasslands . Soil Biol. Biochem . 2005 , 37 , 1707 – 1717 . OpenUrl 74. ↵ Erktan , A. ; Or , D. ; Scheu , S. The physical structure of soil: Determinant and consequence of trophic interactions . Soil Biol. Biochem ., 2020 , 148 , 107876 . OpenUrl 75. ↵ Potapov , M. Multifunctionality of belowground food webs: Resource, size and spatial energy channels . Biol. Rev ., 2022 , 97 , 1691 – 1711 . OpenUrl CrossRef 76. Jílková , V. ; Šebek , O. ; Frouz , J. Mechanisms of pH change in wood ant (Formica polyctena) nests . Pedobiologia 2012 , 55 , 247 – 251 . doi: 10.1016/j.pedobi.2012.04.002 . OpenUrl CrossRef 77. ↵ Castaño-Meneses , G. ; Palacios-Vargas , J.G. ; Delabie , J.H.C. ; Zeppelini , D. ; Mariano , C.S.F. Springtails (Collembola) associated with nests of fungus-growing ants (Formicidae: Myrmicinae: Attini) in southern Bahia, Brazil . Fla. Entomol . 2017 , 100 , 740 – 742 . doi: 10.1653/024.100.0421 . OpenUrl CrossRef 78. ↵ Gwiazdowicz , D.J. Gwiazdowicz , D.J. Mesostigmatid mites (Acari) associated in nests of Formicidae in Poland . In Selected problems of acarological research in forests , Gwiazdowicz , D.J. , Ed.. 2013 . Wydawnictwo Uniwersytetu Przyrodniczego , Poznań, P{oland , 2013 ; pp. 97 – 112 . OpenUrl 79. ↵ Wiśniewski , J. Ergänzung der Deutonymphenbeschreibung von Uropoda (Phaulodinychus) spinosula (Kneissl 1916) (Uropodini, Uropodinae) . Acarologie 1980 , 27 , 15 . OpenUrl 80. ↵ Wiśniewski , J. Für die Fauna Polens neue Uropodina (Acari: Parasitiformes) . Teil II. Fragm. Faun . 1982 , 27 , 143 – 147 . OpenUrl 81. Wiśniewski , J. Studies on a Biological Complex of Factors Regulating Forest Ant Populations . Final Report FG-PO-366; Manuscript , 1983 . 82. ↵ Magura , T. ; Lövei , G.L. Consequences of urban living: Urbanization and ground beetles . Curr. Landsc. Ecol. Rep . 2021 , 6 , 9 – 21 . doi: 10.1007/s40823-020-00060-x . OpenUrl CrossRef 83. ↵ Gwiazdowicz , D.J. Roztocze (Acari, Gamasida) występujące w mrowiskach Formica polyctena Förster na terenie Białowieskiego Parku Narodowego . Parki Nar. Rez. Przyr . 2001 , 20 , 97 – 101 (in Polish) OpenUrl View the discussion thread. Back to top Previous Next Posted August 02, 2025. Download PDF Email Thank you for your interest in spreading the word about bioRxiv. NOTE: Your email address is requested solely to identify you as the sender of this article. Your Email * Your Name * Send To * Enter multiple addresses on separate lines or separate them with commas. You are going to email the following Impact of Biogenic Structures of the Soil-Nesting Ants Lasius niger and Lasius flavus on the Soil Microarthropod Community in Urban Green Spaces Message Subject (Your Name) has forwarded a page to you from bioRxiv Message Body (Your Name) thought you would like to see this page from the bioRxiv website. Your Personal Message CAPTCHA This question is for testing whether or not you are a human visitor and to prevent automated spam submissions. Share Impact of Biogenic Structures of the Soil-Nesting Ants Lasius niger and Lasius flavus on the Soil Microarthropod Community in Urban Green Spaces Maria Sterzyńska , Dariusz J. Gwiazdowicz , Paweł Nicia , Paweł Zadrożny , Gema Trigos-Peral , Mohamed W. Negm bioRxiv 2025.07.31.667896; doi: https://doi.org/10.1101/2025.07.31.667896 Share This Article: Copy Citation Tools Impact of Biogenic Structures of the Soil-Nesting Ants Lasius niger and Lasius flavus on the Soil Microarthropod Community in Urban Green Spaces Maria Sterzyńska , Dariusz J. Gwiazdowicz , Paweł Nicia , Paweł Zadrożny , Gema Trigos-Peral , Mohamed W. Negm bioRxiv 2025.07.31.667896; doi: https://doi.org/10.1101/2025.07.31.667896 Citation Manager Formats BibTeX Bookends EasyBib EndNote (tagged) EndNote 8 (xml) Medlars Mendeley Papers RefWorks Tagged Ref Manager RIS Zotero Tweet Widget Facebook Like Google Plus One Subject Area Ecology Subject Areas All Articles Animal Behavior and Cognition (7618) Biochemistry (17633) Bioengineering (13856) Bioinformatics (41841) Biophysics (21399) Cancer Biology (18529) Cell Biology (25422) Clinical Trials (138) Developmental Biology (13352) Ecology (19860) Epidemiology (2067) Evolutionary Biology (24282) Genetics (15582) Genomics (22462) Immunology (17700) Microbiology (40295) Molecular Biology (17140) Neuroscience (88419) Paleontology (666) Pathology (2823) Pharmacology and Toxicology (4813) Physiology (7632) Plant Biology (15107) Scientific Communication and Education (2042) Synthetic Biology (4284) Systems Biology (9808) Zoology (2267)