Content uploaded by Adrian Purkart
Author content
All content in this area was uploaded by Adrian Purkart on Mar 28, 2023
Content may be subject to copyright.
J. Entomol. Res. Soc., 25(1): 79-90, 2023 Research Article
Doi: 10.51963/jers.v25i1.2183 Online ISSN:2651-3579
Langraf, V., Purkart, A., Petrovičová, K., & Schlarmannová, J. (2023). The community structure of ants in
Hordeum vulgare and grass mixture conditions in the southwestern part of Slovakia. Journal of the
Entomological Research Society, 25(1), 79-90.
Received: January 19, 2022 Accepted: February 19, 2023
The Community Structure of Ants in Hordeum Vulgare and Grass
Mixture Conditions in the Southwestern Part of Slovakia
Vladimír LANGRAF1* Adrián PURKART2 Kornélia PETROVIČOVÁ3
Janka SCHLARMANNOVÁ1**
1 Department of Zoology and Anthropology, Faculty of Natural Sciences, Constantine the
Philosopher University in Nitra, Tr. A. Hlinku 1, Nitra, SLOVAK REPUBLIC
2 Department of Zoology, Faculty of Natural Sciences, Comenius University, Ilkovičova 6,
842 15 Bratislava, SLOVAK REPUBLIC
3 Institute of Plant and Environmental Sciences, Faculty of Agrobiology and Food Resources
Slovak, University of Agriculture in Nitra, Tr. A. Hlinku 2, 94901 Nitra, SLOVAK REPUBLIC
e-mails: 1*langrafvladimir@gmail.com, 2pu rkart .adri an@gmail .com,
3kornelia.petrovicova@gmail.com 1**jschlarmannova@ukf.sk
ORCID IDs:1*0000-0002-3839-3036, 20000-0002-8471-5083
30000-0002-1581-2517, 1**0000-0003-0730-6056
1*Corresponding Authors
ABSTRACT
Ants are an important bioindicative group that plays a signicant role in agroecosystems. As a result of
interspecic competition for food, they can displace native species. The aim of the research was to assess
the inuence of environmental variables (soil pH, soil moisture, potassium, phosphorus and nitrogen) and
the inuence of seasons on the dispersion of ants. Between 2018 and 2020, while investigating dierent
types of crops, we recorded 864 individuals belonging to 9 species and 2 unspecied species (sp.). The
dispersion of ants was aected by moisture, soil pH, phosphorus, potassium and nitrogen. In addition, an
increase in value of the average number of individuals during spring and summer months was conrmed.
We conrmed an increasing number of ant individuals with increasing values of potassium, phosphorus,
nitrogen and soil moisture. A neutral pH of soil is optimal for ants. Our results yielded new information
indicating that agricultural intensication negatively aects ants which are important for the production of
biomass and reduces the number of pests which also aect crop yields.
Keywords: ants, abundance, agrosystems, diversity, eld margins.
80
LANGRAF, V., PURKART, A., PETROVIČOVÁ, K., & SCHLARMANNOVÁ, J.
INTRODUCTION
In a world with a rapidly expanding population of people, there is a growing demand
for food and a simultaneous need for higher environmental sustainability. The area
available for agricultural production is limited, and methods which do not compromise
yields are needed. Insects are one of the largest groups of animals which play a vital
part in the conservation of ecosystems, improve the health of an ecosystem, and
are the critical component in the food web in both terrestrial and aquatic ecosystems
(Courtney, 1994; Brygadyrenko, 2015; Faly, Kolombar, Prokopenko, Pakhomov, &
Brygadyrenko, 2017; Avtaeva, Sukhodolskaya, & Brygadyrenko, 2021). Ants are one of
the most ecologically dominant groups of insects in terrestrial habitats. Their ecological
success can be attributed to the variety and eciency of their foraging habits, eusocial
mode of life, local abundance and the ability to adjust their activity to environmental
changes (Ronque, Fourcassié, & Oliveira, 2018). They are a social insect group that
carry out various roles such as predator, prey, detritivore and herbivore (Diamé, Rey,
Vayssières, Grechi, Chailleux, & Diarra 2018). They vary signicantly, there are almost
14,000 species found widely distributed across the earth (Bolton, 2021). They are
cosmopolitan and exist across several dierent ecosystems, including forests, damp
places, water sources and drylands. Given this presence, it is no wonder that ant
diversity is used as a bioindicator to determine ecosystem and environmental changes
(Gibb, et al, 2020; Oberprieler & Andersen, 2020).
A
nts contribute to various ecosystem services including soil dynamics and nutrient
cycling, they directly aect species composition in animal and plant communities
(Toro, Ribbons, & Pelini, 2012), they represent an important component of agricultural
ecosystems (Oenberg, 2015) and especially semi-natural habitats within agricultural
landscapes (Marshall & Moonen, 2002). They are skilful tillers of soil, dispensers
of seeds and microbial propagules, transmitters of N2-xing bacteria, ecosystem
engineers, fungi growers, waste managers, biotechnologists, pest controllers, soldiers
and reproducers (Benckiser, 2007). Their densities and compositions in agricultural
sites depend on human activities and are predictable in a typical agricultural land-use
mosaic such as arable, fallow, grassland elds and forest sites (Dauber, 2001; Braschler,
2005; Purkart, Kollár, & Goová, 2019). Therefore, many ant species hold desirable
characteristics unshared by most other benecials. They comprise at least one-third of
all insect biomass (Hölldobler & Wilson, 2009). With such abundance, any interaction
derived from this taxon holds a high potential. Most ant species are polyphagous,
cooperative often with polymorphic worker forces, enabling them to deploy a wide range
of prey types. They may exert pressure on several pest species and their life stages.
Their territorial behaviour makes them attack and deter pests that are far beyond the
size of potential prey (Manak, Nordenhem, Bjorklund, Lenoir, & Nordlander, 2013).
Weaver ants (Oecophylla smaragdina and O. longinoda) control more than 50 dierent
pests in 12 dierent crops. They are able to increase farmers’ net income by more than
70% when substituting conventional pesticide regimes (Peng, Christian, & Gibb, 2004;
Peng & Christian, 2005). Ants’ predation makes them prospects for future integrated
pest management strategies in agriculture
(Oenberg, 2014).
81
The Community Structure of Ants in Hordeum Vulgare
The objective of this study was to analyse the community structure of ants in the
conditions of Hordeum vulgare and Grass mixture. In the work, we also analysed the
eects of environmental variables (soil humidity, soil pH, potassium, phosphorus and
nitrogen), which might inuence their abundance and population structure. The results
of our work might inuence for the correct setting of crop management, so that there
is no disturbance to the population of ants due to their importance in agroecosystems,
where they are part of the biomass and also participate in the reduction of crop pests.
MATERIALS AND METHODS
The research took place in the year 2018 to 2020 and we collected ants in two types
of agricultural crops. In the winter crop of Hordeum vulgare, ants were collected from
November to July. In the Grass mixture, ants were collected year-round. These types
of agricultural crops were examined throughout each year, the position of crops in the
elds changed every year (Klimánek, 2008). Crops were grown in a conventional way.
The soil was ploughed three times and turned. Pre-sowing preparation and sowing
were combined. Machines with driven working tools were used in conjunction with a
seed drill. When sowing, it was possible to use seed coulters with an obtuse angle
of penetration into the soil.
Study area
The study area of agricultural crops is located in the geomorphological unit of the
Podunajská pahorkatina - the Danubian upland (the south-western part of Slovakia)
in the cadastral territory of Nitra Fig. 1. The altitude of the monitored area was
approximately 130m above sea level with a brown type of soil. The study area belongs
to a warm arid climate area with mild winters (Table 1).
Figure 1. Map of the study area.
82
LANGRAF, V., PURKART, A., PETROVIČOVÁ, K., & SCHLARMANNOVÁ, J.
Table 1. Average values of temperature and rainfall.
Month Average temperature (°C) Average rainfall (mm)
January −5–5 30
February −3–6 26
March 0–12 35
April 10–20 12
May 15–22 65
June 18–27 77
July 22–29 41
August 20–29 57
September 15–23 64
October 8–15 54
November −3–7 40
December −5–5 55
Collection of samples and application of sprays
We used ve pitfall traps for each site, which were placed in a line at a distance
of 10 metres. A 4% formaldehyde solution to x the material was used. Pitfall traps
were always in the elds and were collected at two-week intervals. The nomenclature
and determination of ants was established according to the work of Seifert (2018).
The insecticide FORCE (Syngenta, Basel, Switzerland), a granular insecticide
in-tended for soil application to control soil pests, was applied to the crops. Insects
were killed through respiratory and tactile poison ingestion. The preparation had a
fast eect and a strong residual (repellent) action against a wide range of soil pests
from the orders of Coleoptera, Aranea and Hymenoptera. The applied dose was
administered uniformly at a concentration of 12–15 kg per ha each year for the duration
of the research. Solinure FX fertiliser (Medilco Hellas S.A., Athens, Greece) containing
chlorides and urea, was applied to the crops and was intended for eld fertility. Due
to its acidifying eect, it contributed to lowering the soil pH.
Measurement of environmental variables
At each pitfall trap location we removed stones and fallen leaves from crops and
sampled the soil to a depth of 15 cm for analysis. Five samples (one from each of
ve sites) were taken from each eld every two weeks over the three years of the
study period. Subsequently, environmental variables (N, P, K, pH, moisture) were
analysed using a soil moisture meter (Rapitest 3 1835, Luster Leaf, Illinois, IL, USA)
and a pH meter (Dexxer PH-03, Luboň, Poland). We thoroughly wetted the broken
up soil with water (ideally distilled or deionised water) to a muddy consistency. We
wiped the meter probe clean with a tissue or paper towel and inserted it into the soil
up to the probe base (7-10 cm). We waited one minute and wrote down the value.
We converted the measured values into units of mg.
Database quality
The data obtained by research has been saved in the Microsoft SQL Server 2017
database program (Express Edition), consisting of frequency tables for collections
and measured environmental variables, (pH, soil moisture, potassium, phosphorus
83
The Community Structure of Ants in Hordeum Vulgare
and nitrogen). The database also consisted of code tables for study sites and their
variables (crops, habitat, locality name, cadastral area, altitude and coordinates of
localities). Matrices for statistical calculations using the Microsoft SQL Server 2017
were programmed.
Statistical analyses
The multivariate analysis (redundancy analysis - RDA) to determine the
dependencies between objects (ants, agricultural crops and soil characteristics) was
used. We tested the statistical signicance of pH, soil moisture, potassium, phosphorus
and nitrogen with the Monte Carlo permutation test in the CANOCO5 program (Ter
Braak & Šmilauer, 2012).
Analysis in the statistical program Statistica Cz. (StatSoft Inc., 2004) focused on
polynomial regression, expressing the relationship between the number of ants and
the values of potassium, phosphorus, nitrogen, pH and soil moisture. The Shapiro-Wilk
W-test determined the normality of data distribution. Based on the violation of the
normality data distribution (p-value = 0.00), we used the nonparametric Friedman test
(ANOVA). It was used to test the dierences in the number of individuals between
the months.
RESULTS
Over a period of three years of research, we found a total of 864 individuals
belonging to 9 species and 2 unspecied species (sp.) in the studied area. Species
of Lasius niger (83.80%) and Tetramorium caespitum (11.11%) had a eudominant
representation of individuals, other species had subdominant to subrecendent
representation (Table 2).
Table 2. Distribution of the ants in the agricultural crops during the years 2018 - 2020.
Species Grass mixture Hordeum vulgare ∑ ind.
Formica cunicularia Latreille, 1798 411 15
Formica rubarbis Fabricius, 1793 0 2 2
Formica sp. 0 1 1
Lasius alienus Förster, 1850 0 1 1
Lasius niger (Linné, 1758) 305 419 724
Lasius sp. 3 3 6
Lasius umbratus (Nylander, 1846) 2 11 13
Myrmica sabuleti Meinert, 1861 0 4 4
Polyergus rufescens (Latreille, 1798) 1 0 1
Solenopsis fugax Latreille, 1798 1 0 1
Tetramorium caespitum Santschi, 1927 1 95 96
∑ individuals 317 547 864
Multivariate analysis of the ants between the years 2018 and 2020 was determined
using the redundancy analysis (RDA, SD = 1.40 on the rst ordination axis). We
observed the relationship between ants and environmental variables (pH of the soil, soil
moisture, potassium, phosphorus and nitrogen). The values of the explained variability
of taxonomic data were 50.9% on the rst ordination axis and 54.8% on the second
84
LANGRAF, V., PURKART, A., PETROVIČOVÁ, K., & SCHLARMANNOVÁ, J.
ordination axis. The cumulative variability of the species set explained by environment
variables was represented in the rst ordination axis 88.9% and in the 2nd axis 95.7%.
Using the Monte Carlo permutation test, we identied a statistically signicant eect of
soil moisture (p = 0.0088, F(1.0276) = 2.0021, df = 5), soil pH (p = 0.0508, F(1.1183)
= 1.9297,df = 5), phosphorus (p = 0.0466, F(1.1952) = 2.0805, df = 5), potassium (p =
0.0328, F(1.7145) = 1.9620, df = 5) and nitrogen (p = 0.0490, F(1.7006) = 2.1005, df =
5) on the structure of arthropods. The selected variables were not mutually correlated
with the maximum value of the ination factor = 4.3243. The ordination graph (triplot)
contained ants ordered into one cluster (Fig. 2). The rst cluster (I) consisted of ants
correlated with phosphorus (mg) and moisture. The Formica rubarbis species has
links to potassium (mg) and soil pH. Polyergus rufescens correlated with nitrogen
(mg). Solenopsis fugax was not aected by environmental variables.
Figure 2. RDA analysis of ants with environmental variables.
The normality data distribution (number of individuals) was violated (p-value =
0.00). Based on that, a nonparametric Friedman test (ANOVA) was used to conrm
the statistically signicant dierence (p-value = 0.04670, F (2.27) = 2.83040, df = 3)
(Fig. 3) of individuals between months and crops of the Hordeum vulgare and Grass
mixture. The results showed an increase in the average value of individuals in June
- August in the crops Hordeum vulgare. Under Grass mixture conditions, the number
of individuals increased from March to June and decreased in the following months.
The number of individual ants was processed using polynomial regression.
Using the regression model, we expressed the relationship (correlation) between
the number of individuals of ants and potassium (mg), phosphorus (mg), nitrogen
(mg), pH and humidity (%). The correlation coecient value was high for the number
of individuals and pH (r = 0.8140) (Fig. 4, A), potassium (r = 0.9012) (Fig. 4, B),
phosphorus (r = 0.8905) (Fig. 4, C), nitrogen (r = 0.7981) (Fig. 4, D) and moisture (r
= 0.881) (Fig. 4, E), which indicated a strong relationship. The reliability coecient
for the pH r2 = 0.6899 indicated the capture of 68% variability, potassium r2 = 0.6908
(69% variability), phosphorus r2 = 0.7504 (75% variability), nitrogen r2 = 0.7145
(71% variability) and moisture r2 = 0.7384 (73% variability). The overall suitability of
the regression model is statistically signicant in all cases: pH (p-value = 0.0015),
85
The Community Structure of Ants in Hordeum Vulgare
potassium (p-value = 0.0428), phosphorus (p-value = 0.0298), nitrogen (p-value =
0.0248) and moisture (p-value = 0.0118). The results showed that increasing values
of potassium, phosphorus, nitrogen and soil humidity also increased the number of
ant individuals. The ideal value for ants was 16 - 22 mg/kg potassium, 1.3 - 1.8 mg/
kg phosphorus, 16 - 22 mg/kg nitrogen, 7 pH and 14 – 22 % for moisture.
Figure 3. Friedman test (ANOVA) dierence in the number of individuals between month and crops.
Figure 4. Polynomial regression model potassium, phosphorus, nitrogen, pH and moisture on the number
of individuals of the ants.
DISCUSSION
Ants living in agricultural landscapes have a wider tolerance than ants from natural
habitats. They can achieve high local density due to the inuence of agriculture and eld
margins support the most diverse community of ants (Bote & Romero, 2012; Oliveira et
al, 2012; Magura, Ferrante, & Lövei 2020). We recorded that the ant community was
86
LANGRAF, V., PURKART, A., PETROVIČOVÁ, K., & SCHLARMANNOVÁ, J.
dominated by species Lasius niger and Tetramorium caespitum. The great abundance
of ants aects the maintenance of the natural balance and substance cycle of the
biogenic elements in ecosystems such as carbon, nitrogen, sulphur and phosphorus.
The dominance of Hymenoptera (Formicidae) and Coleoptera has been indicated as
a general trait of ground-dwelling assemblages (Miranda, Piñero, & Megías, 2007;
Lenoir & Lennartsson, 2010; Pardee & Philpott, 2011). Their activities accelerated
the decomposition of plant residues, aerated the soil and improved soil structure
and quality (Holecová, Lukáš, & Harakaľová, 2003; Dieng, Ndiaye, & Taylor, 2016).
The dominant representation of the ants (Formicidae) and Coleoptera taxon among
epigeic arthropods in the conditions of integrated farming and ecological farming was
also recorded by Porhajašová Noskovič, Rakovská, Babošová, & Čeryová (2015);
Porhajašová, Babošová, Noskovič, & Ondrišík (2018).
Exploiting biodiversity on ecosystem service provision is a goal of contemporary
agriculture, although relationships between diversity and ecosystem services remain
largely unexplored for innovative practices (Kalivoda, Petrovič,& Kürthy, 2010; Finney
& Kaye, 2016; Griths et al, 2000; Špulerová, Petrovič, Mederly, Mojses, & Izakovičová
2018; Dobrovodska, Kanka, & David, 2019). Ants play an irreplaceable role in the
decomposition of organic matter, in the cycle of biogenic elements of carbon, nitrogen,
sulphur, phosphorus, in transformation and degradation of waste and toxic substances,
and their presence is irreplaceable (Fazekašová & Bobuľovská, 2012). Using the
multivariate model, we demonstrated the inuence of environmental variables (pH
of the soil, soil moisture, potassium, phosphorus, nitrogen) on the abundance of
ants. Thus, our results agreed with the results of (Attwood, Maron, House, & Zammit
(2008)), who observed a change of abundance of ants with increasing land use.
Biodiversity loss as a consequence of agricultural intensication can lead to reductions
in agroecosystem functions and services. Increasing crop diversity through rotation
may alleviate these negative consequences by restoring positive interactions. The
impact of ants is an important component of the strategy leading to the sustainability
of the soil ecosystem. The diversity of ants, including its abundance in soil, depends
on the abiotic and biotic factors that are typical of the biotope (Zak, Holmes, White,
Peacock, & Tilman, 2003; Tiemann, Grandy, Atkinson, Spiotta, & McDaniel, 2015).
Arthropod abundance from month to month is usually interpreted as being related
to uctuations in climatic factors (such as temperature, precipitation and day length)
(Lionello, Rizzoli, & Boscolo, 2006). The number of ants in March and June was
higher from the number of ants captured in July and September in Grass mixture
conditions. In the Hordeum vulgare crop we have seen a steady increase. Simão,
Carretero, Amaral, Soares, & Mateos (2015) also conrmed dierences in the
number of ants aected by dierent weather during the seasons. Andrew, Roberts,
Hill (2012) have suggested that precipitation is more inuential on ant diversity at
high temperatures than at low temperatures. Greenberg & McGrane (1996); Majeed,
Rana, Azevedo, Elmo, & Nargis (2020) conrmed a seasonal trend for the abundance
of arthropod groups. It is established that environmental variables and the inuence
of biogeographic factors account for uctuations in species abundance. Climatic
87
The Community Structure of Ants in Hordeum Vulgare
conditions during the months impact the biodiversity of ant species (Garcia, Cabeza,
Rahbek, & Araújo, 2014; Williams & Newbold, 2020). In our study, we conrmed
with the help of regression models a strong relationship between the environmental
variables potassium (mg), phosphorus (mg), nitrogen (mg), pH, humidity (%) and
the abundance of ants. For agricultural management, understanding how species’
behaviour varies with environmental variables is imperative in ensuring food security
in the future. In addition, ants’ predation makes them prospects for future integrated
pest management strategies in agriculture (Oenberg, 2014). Ants mineralize nutrients,
form soil aggregates, and disperse seeds, are signicant and necessary for decorous
ecosystem functioning and sustainability (Del Toro, Ribbons, & Pelini, 2012; Pfeier,
Mezger, & Dyckmans 2013).
CONCLUSION
Our results have provided new knowledge about the preference of ants in the
conditions of Hordeum vulgare and Grass mixture in central Europe. The dispersion
of ants was inuenced by soil moisture, soil pH, phosphorus, potassium and nitrogen.
We conrmed an increase in the average number of individuals during spring and
summer months. The ants had a strong correlation with soil moisture (%), soil pH,
phosphorus (mg/kg), potassium (mg/kg) and nitrogen (mg/kg). With increasing
values of potassium, phosphorus, nitrogen and moisture, the number of individuals
also increased. We conrmed that the optimal soil pH value was neutral. A practical,
workable approach should be used to preserve the current ant population, achieve
sustainable levels of biodiversity, key species to develop conservation and agricultural
management strategies. This is of particular importance to those who may face
pressure from pest species threatening crop yields. This study can be helpful in the
planning of conservation programs as well as provide information to farmers to initiate
integrated pest management strategies.
ACKNOWLEDGEMENT
This research was supported by the grants VEGA 2/0022/23 and KEGA No.
002UKF-4/2022 Metaanalyzes in biology and ecology (databases and statistical data
analysis).
REFERENCES
Andrew, N. R., Roberts, I. R., & Hill, S. J. (2012). Insect herbivory along environmental gradients. Open
Journal of Ecology, 2(4), 202-213.
Attwood, S. J., Maron, M., House, A. P. N., & Zammit, C. (2008). Do arthropod assemblages display
globally consistent responses to intensied agricultural land use and management? Global Ecology
and Biogeogaphy, 17(5), 585-599.
Avtaeva, T. A., Sukhodolskaya, R. A., & Brygadyrenko, V. V. (2021). Modeling the bioclimating range of
Pterostichus melanarius (Coleoptera, Carabidae) in conditions of global climate change. Biosystems
Diversity, 29(2), 140-150.
88
LANGRAF, V., PURKART, A., PETROVIČOVÁ, K., & SCHLARMANNOVÁ, J.
Benckiser, G. (2007). Principles behind order and sustainability in natural successions and agriculture. In
G., Benckiser, S., Schnell (Eds.). Biodiversity in agricultural production systems (pp. 349-383). Boca
Raton (USA): Taylor & Francis.
Bolton, B. (2012, February 9). An online catalog of the ants of the world. Retrieved from https://antcat.org.
Bote, P. J., & Romero, A. (2012). Epigeic soil arthropod abundance under dierent agricultural land uses.
Spanish Journal of Agricultural Research, 10(1), 55-61.
Braschler, B. M. (2005). Eects of experimental small-scale grassland fragmentation on the population
dynamics on invertebrates. Ph.D. thesis, Switzerland, University of Basel.
Brygadyrenko, V. V. (2015). Community structure of litter invertebrates of forest belt ecosystems in the
Ukrainian steppe zone. International Journal of Environmental Research, 9(4), 1183-1192.
Carvalho, R. L., Andersen, A. N., Anjos, D. V., Pacheco, R., Chagas, L., & Vasconcelos, H. L. (2020).
Understanding what bioindicators are actually indicating: Linking disturbance responses to ecological
traits of dung beetles and ants. Ecological Indicators, 108.
Courtney, G. W. (1994). Biosystematics of the Nymphomyiidae (Insecta: Diptera): Life History, Morphology,
and Phylogenetic Relationships. Smithsonian Institution Press, Washington, D.C.
Dauber J. (2001). Ant communities of an agricultural landscape: Relationships to landscape structure and
land-use management. Ph.D. thesis, Justus Liebig-University of Giessen, Germany.
Del Toro, I., Ribbons, R. R., & Pelini, S. L. (2012). The little things that run the world revisited: a review
of ant-mediated ecosystem services and disservices (Hymenoptera: Formicidae). Myrmecological
News, 17, 133-146.
Diamé, L., Rey, J. Y., Vayssières, J. F., Grechi, I., Chailleux, A., & Diarra, K. (2018). Ants: Major functional
elements in fruit agro-ecosystems and biological control agents. Sustainability, 10(1), 23.
Dieng, M. M., Ndiaye, A. B., Ba, C. T., & Taylor, B. (2016). Les fourmis (Hym., Formicidae) de l’enclos
d’acclimatation de Katane de la réserve de faune du Ferlo nord (Sénégal). International Journal of
Biological and Chemical Sciences, 10(4), 1626-1636.
Doblas-Miranda, E., Sánchez-Piñero, F., & González-Megías, A. (2007). Soil macroinvertebrate fauna of
a Mediterranean arid system: Composition and temporal changes in the assemblage. Soil Biology
and Biochemistry, 39(8), 1916-1925.
Dobrovodska, M., Kanka, R., & David, S. (2019). Assessment of the biocultural value of traditional
agricultural landscape on a plot-by-plot level: case studies from Slovakia. Biodiversity and
Conservation, 28(10), 2615-2645.
Faly, L. I., Kolombar, T. M., Prokopenko, E. V., Pakhomov, O. Y., & Brygadyrenko, V. V. (2017). Structure
of litter macrofauna communities in poplar plantations in an urban ecosystem in Ukraine. Biosystems
Diversity, 25(1), 29-38.
Fazekašová, D. & Bobuľovská, L. (2012). Soil organisms as an Indicator of Quality and Environmental
Stress in the Soil Ecosystem. Environment, 46(2), 103-106.
Finney, D. M. & Kaye, J. P. (2016). Functional diversity in cover crop polycultures increases multifunction-
ality of an agricultural system. Journal of Applied Ecology, 54(2), 509-517.
Garcia, R. A., Cabeza, M., Rahbek, C., & Araújo, M. B. (2014). Multiple dimensions of climate change and
their implications for biodiversity. Science, 344(6183), 486-496.
Gibb, H., Sanders, N. J., Dunn, R. R., Watson, S., Photakis, M., Abril, S., Andersen, A. N., Angulo,
E., Armbrecht, I., Arnan, X., Baccaro, F. B., Bishop, T. R., Boulay, R., Castracani, C., Del Toro, I.,
Delsinne, T., Diaz, M., Donoso, D. A., Enríquez, M. L., Fayle, T. M., Feener, D. H., Fitzpatrick, M. C.,
Gómez, C., Grasso, D. A, Groc, S., Heterick, B., Homann, B. D., Lach, L., Lattke, J., Leponce, M.,
Lessard, J. P., Longino, J., Lucky, A., Majer, J., Menke, S. B,, Mezger, D., Mori, A., Munyai, T. C.,
Paknia, O., Pearce-Duvet, J., Pfeier, M., Philpott, S. M., de Souza, J. L. P., Tista, M., Vasconcelos, H.
L., Vonshak, M., & Parr, C. L. (2015). Climate mediates the eects of disturbance on ant assemblage
structure. Proceedings of the Royal Society B Biological Sciences, 282(1808), 20150418.
89
The Community Structure of Ants in Hordeum Vulgare
Griths, B. S., Ritz, K., Bardgett, R. D., Cook, R., Christensen, S., Ekelund, F., Sørensen, S.J., Bååth, E.,
Bloem, J., De Ruiter, P. C., Dolng, J., & Nicolardot, B. (2000). Ecosystem response of pasture soil
communities to fumigation-induced microbial diversity reductions: an examination of the biodiversity–
ecosystem function relationship. Oikos, 90(2), 279-294.
Greenberg, C. H. & McGrane, A. (1996). A comparison of relative abundance and biomass of
ground-dwelling arthropods under dierent forest management practices. Forest Ecology and
Management, 89(1-3), 31-41.
Holecová, M., Lukáš, J., & Harakaľová, E. (2003). Mravce (Hymenoptera, Formicidae) dubovo-hrabových
lesov v okolí Bratislavy (JZ Slovensko). Folia faunistica Slovaca, 8, 63-69.
Hölldobler, B. & Wilson, E. O. (2009). The superorganism. New York: W.W. Norton & Company.
Kalivoda, H., Petrovič, F., & Kürthy, A. (2010). Inuence of the landscape structure on the buttery
(Lepidoptera, Hesperioidea and Papilionoidea) and bird (Aves) taxocoenoses in Vel’ké Leváre (SW
Slovakia). Ekologia (Bratislava), 29(4), 337-359.
Klimánek, M. (2008). Geoinformation systems tutorials for exercises in the ArcGIS system, Mendel
University in Brno.
Lenoir, L. & Lennartsson, T. (2010). Eects of timing of grazing on arthropod communities in semi-natural
grasslands. Journal of Insect Science, 10, 60.
Lionello, P., Malanotte-Rizzoli, P., & Boscolo, R. (Eds). (2006). Mediterranean Climate Variability. (1st
ed.). Elsevier Science.
Magura, T., Ferrante, M., & Lövei, L. G. (2020). Only habitat specialists become smaller with advancing
urbanization. Global Ecology and Biogeogaphy, 29(11), 1978-1987.
Majeed, W., Rana, N., de Azevedo, K., Elmo, B., & Nargis, S. (2020). Seasonality and Climatic Factors
Aect Diversity and Distribution of Arthropods Around Wetlands. Pakistan Journal of Zoology, 52(6),
2135-2144.
Manak, V., Nordenhem, H., Bjorklund, N., Lenoir, L., & Nordlander, G. (2013). Ants protect conifer seedlings
from feeding damage by the pine weevil Hylobius abietis. Agricultural and Forest Entomology, 15(1),
98-105.
Marshall, E. J. P. & Moonen, A. C. (2002). Field margins in northern Europe: their functions interactions
with agriculture. Agriculture, Ecosystems & Environment, 89(1-2), 5-21.
Microsoft SQL Server (2017). (RTM) - 14.0.1000.169 (X64) Aug 22 2017 17:04:49 Copyright (C) 2017
Microsoft Corporation Express Edition (64-bit) on Windows 10 Home 10.0 <X64> (Build 18362:).
Oberprieler, S. K. & Andersen, A. N. (2020). The importance of sampling intensity when assessing
ecosystem restoration: Ants as bioindicators in northern Australia. Restoration Ecology, 28(4),
737-741.
Oenberg, J. (2014). Pest repelling properties of ant pheromones. IOB-CWPRS Bulletin, 99, 173-176.
Oenberg, J. (2015). Ants as tools in sustainable agriculture. Journal of Applied Ecology, 52(5), 1197-1205.
Oliveira, R. F., Almeida, L. C., Souza, D. R., Munhae, C. B., Bueno, O. C., & Morini, S. C. (2012). Ant
diversity (Hymenoptera: Formicidae) and predation by ants on the dierent stages of the sugarcane
borer life cycle Diatraea saccharalis (Lepidoptera: Crambidae). European Journal of Entomology,
109(3), 381-387.
Pardee, G. L. & Philpott, S. M. (2011). Cascading indirect eects in a coee agroecosystem: eects
of parasitic phorid ies on ants and the coee berry borer in a high-shade and low-shade habitat.
Environmental Entomology, 40(3), 581-588.
Peng, R. K. & Christian, K. (2005). Integrated pest management in mango orchards in the Northern
Territory Australia, using the weaver ant, Oecophylla smaragdina, (Hymenoptera: Formicidae) as a
key element. International Journal of Pest Management, 51(2), 149-155.
Peng, R. K., Christian, K., & Gibb, K. (2004). Implementing ant technology in commercial cashew
plantations and continuation of transplanted green ant colony monitoring. Canberra (Australia): Rural
Industries Research and Development Corporation.
90
LANGRAF, V., PURKART, A., PETROVIČOVÁ, K., & SCHLARMANNOVÁ, J.
Pfeier, M., Mezger, D., & Dyckmans, J. (2013). Trophic ecology of tropical leaf litter ants (Hymenoptera:
Formicidae) - a stable isotope study in four types of Bornean rain forest. Myrmecological News, 19,
31-41.
Porhajašová, J., Noskovič, J., Rakovská, A., Babošová, M., & Čeryová, T. (2015). Biodiversity and
Dynamics of Occurence of Epigeic Groups in Dierent Types of Farming. Acta Horticulturae et
Regiotecturae, 1, 5-10.
Porhajašová, J., Babošová, M., Noskovič, J., & Ondrišík, P. (2018). Long-Term Developments and
Biodiversity in Carabid and Staphylinid (Coleoptera: Carabidae and Staphylinidae) Fauna during the
Application of Organic Fertilizers under Agroecosystem Conditions. Polish Journal of Environmental
Studies, 27(5), 2229-2235.
Purkart, A., Kollár, J., & Goová, K. (2019). Fauna of Ants (Hymenoptera: Formicidae) of Selected Sand
Habitats in Podunajsko Region. Naturae tutela, 23(1), 101-111.
Ronque, U. V. M., Fourcassié, R. V., & Oliveira. P. S. (2018). Ecology and eld biology of two dominant
Camponotus ants (Hymenoptera: Formicidae) in the Brazilian savannah. Journal of Natural History,
52(3-4), 237-252.
Simão, F., Carretero, M. A., Amaral, M. J., Soares, A. M. V. M., & Mateos, E. (2015). Composition and
seasonal variation of epigeic arthropods in eld margins of NW Portugal. Turkish Journal of
Zoology, 39(3), 404-411.
Statsoft, INC. Statistica Cz [Softwarový systém na analýzu dat]. (2004). Www.StatSoft.Cz.
Seifert, B. (2018). The Ants of Central and North Europe. Germany: Lutra Verlags und Vertiebsgesells-
chaft, Tauer.
Špulerová, J., Petrovič, F., Mederly, P., Mojses, M., & Izakovičová, Z. (2018). Contribution of Traditional
Farming to Ecosystem Services Provision: Case Studies from Slovakia. Land, 7(2), 74.
Ter Braak, C. J. F. & Šmilauer, P. (2012). Canoco reference manual and user’s guide: software for
ordination, version 5.0, Ithaca USA, Micro-computer Power.
Tiemann, L. K., Grandy, A. S., Atkinson, E. E., Marin-Spiotta, E., & McDaniel, M. D. (2015). Crop rotational
diversity enhances belowground communities and functions in a agroecosystem. Ecology Letters,
18(8), 761-771.
Williams, J. J. & Newbold, T. (2020). Local climatic changes aect biodiversity responses to land use: A
review. Diversity and Distributions, 26(1), 76-92.
Zak, D. R., Holmes, W. E., White, D. C., Peacock, A. D., & Tilman, D. (2003). Plant diversity, soil microbial
communities, and ecosystem function: are there any links? Ecology, 84(8), 2042-2050.