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. 2025 Oct 3;40(10):188. doi: 10.1007/s10980-025-02207-9

How does landscape composition and configuration affect dung beetle communities in Eastern Austrian agricultural landscapes?

Raja Imran Hussain 1,✉, Benjamin Knittl 1, Christian H Schulze 2, Thomas Frank 1
PMCID: PMC12494615  PMID: 41048254

Abstract

Context

Understanding the effects of landscape composition on biodiversity is crucial, especially in human-dominated agricultural landscapes. This study focuses on dung beetles, an ecologically significant group, to explore how landscape composition and configuration influences species richness, abundance and community structure of dung beetles in Eastern Austrian agricultural areas.

Objectives

The primary objective of this study is to investigate the relationship between landscape composition and dung beetle communities. Specifically, we aim to determine how different habitat types within agricultural landscapes affect dung beetle species richness, abundance and community structure.

Methods

We sampled dung beetles across 14 study landscapes, each with a diameter of 1 km, varying from homogenous landscapes dominated by annual crops to highly heterogeneous landscapes with diverse habitats such as woodlands, hedgerows, and set-aside land (areas left fallow or uncultivated). The study design focused on comparing dung beetle communities across these varying landscape compositions.

Results

Our results reveal that dung beetle species richness is positively correlated with woodland cover, set-aside land and hedgerow length, while beetle abundance is associated with woodland cover and hedgerow length. Additionally, dung beetle communities were highly nested, with communities from landscapes with less woodland cover nested within those from landscapes with higher woodland cover. This underscores the importance of heterogeneously structured landscapes, such as woodlands, for maintaining diverse dung beetle communities. These findings highlight that a decline in structural diversity, often caused by agricultural intensification, likely reduces the ecosystem services provided by dung beetles.

Conclusions

The study’s findings emphasize the significance of maintaining landscape structural diversity to support dung beetle communities and their associated ecosystem services. Recommendations for landscape management and planning include promoting heterogeneous landscapes with set-aside land to enhance biodiversity and ecosystem functioning in agricultural areas.

Keywords: Coprophagous beetles, Landscape heterogeneity, Set-aside land, Species richness, Species composition

Introduction

Agricultural landscapes are some of the most heavily modified ecosystems by humans. The intensification of agriculture has caused a significant decline in biodiversity (Weibull et al. 2003; Shah et al. 2023; Shi et al. 2024). This loss of biodiversity is concerning because many species provide essential ecosystem services, such as pollination, pest control, and nutrient cycling (Albrecht et al. 2012; Blaauw and Isaacs 2014; Tschumi et al. 2016; Bürgler et al. 2024). To maintain biodiversity in these areas, habitat heterogeneity and the presence of semi-natural habitats are crucial (Haaland et al. 2011; Priyadarshana et al. 2024). However, agricultural intensification, which involves the increased use of fertilizers, pesticides, and monocultures, has simplified habitats and reduced farmland biodiversity (Geiger et al. 2010; Tscharntke et al. 2012; Hussain et al. 2021, 2023). Understanding how landscape composition affects biodiversity is therefore essential for developing effective conservation strategies (Landis 2017; Martin et al. 2019; Shah et al. 2023). Further, landscape configurations, such as the spatial arrangement of habitat patches, also play a crucial role in influencing arthropod communities in agricultural landscapes (Rischen et al. 2023). Landscape configuration can affect the movement, survival, and reproduction of arthropods by altering habitat connectivity and the availability of resources (Fahrig et al. 2011; Duflot et al. 2017).

Dung beetles (Scarabaeidae) are an important group of insects that play a key role in ecosystems. They contribute to soil fertilization, nutrient cycling, seed dispersal, and pest control (Nichols et al. 2008; Torabian et al. 2024). Because they are sensitive to environmental changes, dung beetles are often used as bioindicators of landscape health (Nichols et al. 2007; Gardner et al. 2008). Previous research has shown that land-use intensity negatively impacts dung beetle diversity across different ecosystems, from forests to agroforestry systems and annual crops (Shahabuddin et al. 2005; Harvey et al. 2006Shahabuddin et al., 2010). In agricultural landscapes, woodlands and set-aside lands have been found to support higher diversity in arthropod communities (Drapela et al. 2008; Querner et al. 2013), including dung beetles (Kessler et al. 2011; Schernhammer 2020). Set-aside land refers to areas of agricultural land that are left fallow or uncultivated to promote biodiversity and ecosystem health. These habitats provide important resources, such as food and breeding sites, and act as refuges from the disturbances caused by farming activities (Jeanneret et al. 2003; Woodcock et al. 2005).

This study focuses on dung beetle communities in Eastern Austrian agricultural landscapes, which range from simple, homogeneous croplands to complex, heterogeneous landscapes with diverse habitats. We predict that set-aside lands will have a positive effect on dung beetle species richness and abundance by providing critical refuge, diverse microhabitats, and reduced agrochemical exposure, supporting ecological functions and biodiversity persistence (Babu 2018; Escudero et al. 2025). Additionally, we expect that landscapes with higher structural complexity will support more diverse dung beetle communities (Martin et al. 2019). In contrast, intensively managed agricultural landscapes are likely to have simpler communities that are subsets of the richer communities found in more complex landscapes. These predictions are based on the idea that woodlands and set-aside lands habitats provide critical resources and favorable conditions for dung beetles, which are necessary for their survival and reproduction (Frank et al. 2017; Barretto et al. 2020).

Methods

Study area and study sites

Dung beetle communities were sampled at 14 study sites in the eastern part of Lower Austria and in the northern part of Burgenland, Austria (Figure 1). The study sites in Lower Austria were located around the villages of Deutsch-Haslau, Schönabrunn and Prellenkirchen, in Burgenland around Edelstal. The entire study area is part of the Vienna Basin, bounded by the hills Spitzerberg and Hundsheimer Berge in the north and by the Pannonian Basin in the southeast. The main soil type in our study region is black soil (http://www.bodenkarte.at/). The main field crops were cereals followed by maize, rape, sugar beet, potatoes, and wine. The mean annual temperature of our study area is 10.0 °C, the mean annual precipitation is 637 mm (measured at Hainburg; http://de.climate-data.org/location/159305/).

Figure 1.

Figure 1.

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Maps showing (a) the locations of the study landscapes in eastern Austria and (b) the arrangement of the 14 sampled landscapes (circles; major rivers (pale grey line), major roads (black lines), villages (hatched areas) and all larger woodlands (dark grey areas)). Additionally, (c) the spatial distribution of the 9 dung baits (grey squares) placed at the center and at radii of 150 m and 350 m within each study landscape is illustrated.

All 14 study sites were situated at 130−190 m a.s.l and represented “circular landscapes” (subsequently termed “study landscapes”) with a diameter of 1 km (Fig. 1b-c). These sampled study landscapes are a subset of 29 randomly selected landscapes used in the study by Drapela et al. (2008) and represent a gradient from simple landscapes predominantly characterized by crops (mainly cereals and maize) to landscapes with a high structural complexity and habitat diversity. Structural complexity in these landscapes is characterized by factors such as the mean size of crop fields, the connectivity between habitat patches, and the edge densities of these patches. Landscapes with higher structural complexity typically have smaller crop fields, higher connectivity, and greater edge densities, providing more diverse microhabitats and resources for dung beetles. The following landscape variables were measured and used for the analysis in the present study: set-aside land (areas left fallow or uncultivated), hedgerow length, woodland cover (including both continuous forests and tree lines above a certain threshold, i.e. 0.5 hectares for forests and 5 meters for tree lines), annual crop cover (all in % of total area), and structural landscape diversity (Appendix Table A.1). Structural landscape diversity was quantified using two measures: the mean perimeter area ratio (MPAR) of all land cover patches, which indicates landscape-level geometric complexity or fragmentation, and the number of shapes characterizing points (NSCP) index, which represents landscape complexity (Moser et al. 2002). Land cover types were measured using digital orthophotos and satellite imagery analyzed via Geographic Information Systems (GIS).

Table 1.

Results of GLMs testing for effects of NSCP, set-aside land, woodland cover and hedgerow length on estimated species richness and abundance of dung beetles

Variables B SE 95% CI Wald Chi- p
lower Upper Square
Species richness
 (Intercept) 15.39 9.54 − 3.32 34.09 2.60 0.107
 log (NSCP) − 4.01 2.23 − 8.43 0.32 3.31 0.069
 arcsin (set-aside land) 30.30 10.76 9.22 51.39 7.93 0.005
 arcsin (woodland cover) 47.39 10.45 26.90 67.88 20.55 <0.001
 log (hedgerow length + 1) 2.08 0.90 0.32 3.84 5.38 0.020
Abundance
 (Intercept) − 76.831 144.34 − 359.73 206.07 0.28 0.595
 log (NSCP) − 3.77 33.73 − 69.87 62.34 0.01 0.911
 arcsin (set-aside land) − 90.70 162.70 − 409.60 228.19 0.31 0.577
 arcsin (woodland cover) 418.15 158.12 108.23 728.07 6.99 0.008
 log (hedgerow length + 1) 29.06 13.57 2.46 55.66 4.59 0.032
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Sampling and identification of dung beetles

To account for seasonality, dung beetles were sampled in all landscapes four times in May, July, September and October 2007, respectively. During all sampling periods a total of nine dung baits (each consisting of 250 g of fresh horse dung) were exposed in each landscape in similar habitats for three days. At each study landscape one dung bait was placed in its centre, while the others were located at a radius of 150 m (4 traps) and 350 m (4 traps) around it. This spatial design of trapping should enable to sample the majority of dung beetle species of the study landscapes. The used dung was always from the same two horses; none of those received any medical treatment during the whole time, which potentially could have affected the results (e.g. Römbke et al. 2007). While the landscape variables were measured within a 500 m radius based on the center dung bait, we acknowledge that this may not fully capture the surrounding landscape effects for all 9 baits.

Baits were re-collected after three days from the field. Additionally, the soil surface below the baits was collected to a depth of 5–10 cm to include beetles feeding on dung particles buried into the soil beneath the dung pats. During the four sampling periods (May, July, September, October) dung pats were always placed at the same positions. To assure that dung baits can be re-located, locations of dung pats were marked with coloured wooden sticks. Therefore, only 3–4 dung baits out of 126 (2–3%) were lost during each sampling period. To extract the beetles from the dung and the collected soil, a Berlese funnel was used (Southwood 1978). Two 500 W halogen headlights and a 200 W infrared lamp were used to create a gradient of decreasing dung humidity towards the bottom of the dung. Trying to escape the decreasing humidity, beetles crawl deeper into the funnels and fall into jars filled with a killing agent (70% ethanol). To avoid the escape of beetles, the top of the funnel was closed with gauze. To validate this method, dried out dung was investigated for remaining beetles, which demonstrated that nearly 100% of the beetles were successfully extracted from the dung. All caught beetles were preserved in 70% ethanol. In this study only coprophagous beetles belonging to the family Scarabaeidae were considered. Beetles were identified according to Bunalski (1999) and Klausnitzer (1994a, b a, b) and by using a reference collection based at the Department of Botany and Biodiversity Research, University of Vienna.

Data analysis

To estimate the total species richness for each landscape, we used the Jackknife 2 (Jack2) estimator, which is known for its precision and robustness in richness estimation (Walther and Moore 2005). Both species richness (estimated by Jack2) and total beetle abundance were tested for normality using the Kolmogorov-Smirnov test, and neither deviated significantly from a normal distribution (p > 0.20). Therefore, we applied generalized linear models (GLMs) with a normal error distribution and an identity link function to assess the effects of landscape variables on dung beetle richness and abundance. Generalized linear models (GLMs) were constructed using all possible combinations of these variables. The significance of each term was tested through the deviance that the removal of the term adds to the model. Akaike’s Information Criterion (AIC) was used to select the best models, with the lowest AIC indicating the best fit. The effect of MPAR (Mean Perimeter Area Ratio) on dung beetle species richness and abundance was statistically tested using generalized linear models (GLMs), with MPAR included as an independent variable.

Due to a significant negative correlation between crop cover and woodland cover (r = − 0.81, p < 0.001), these variables were not included together in a single model to avoid multicollinearity. Instead, separate models were constructed for each variable.

To analyze the nestedness of species communities, we calculated Monte Carlo-derived probabilities for presence-absence matrices being random. Nestedness was tested by comparing the system temperatures of these matrices with the average temperature of 1000 randomized matrices using the binmatnest algorithm (Rodríguez-Gironés and Santamaría 2006). Spearman rank correlations were calculated to test for relationships between the ranking of sampled landscapes in the presence-absence matrix packed into a state of maximum nestedness and all landscape variables. Subsequently, false discovery rate (FDR) transformations were applied to correct for bias caused by multiple testing (Pike 2011).

Species composition similarity between the 14 study sites was quantified using Bray-Curtis similarities, calculated from square-root transformed abundance data to reduce the influence of dominant species (Clarke and Warwick 2001). These similarity relationships were visualized using non-metric multidimensional scaling (NMDS) ordination. A stress value of < 0.20 was considered to adequately represent the similarity relationships (Clarke 1993). We performed Spearman rank correlations to evaluate the effects of landscape structure on species composition by testing the relationships between NMDS axis scores and landscape variables. This approach allowed us to identify significant correlations between species composition and landscape variables, which are displayed in Figure 3. The landscape variables included in the analysis were annual crops, set-aside land, woodland cover and hedgerow length. All statistical analysis were performed in Statistica version 7.1.

Figure 3.

Figure 3.

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NMDS plot based on Bray-Curtis similarities (square-root transformed abundances) of dung beetles in 14 landscapes. Dimensions 1 and 2 are shown. Pie charts at the plot centers show the percentage cover of crop, set-aside land, woodland, and other habitats. Dimension 1 values were significantly correlated with woodland and crop cover.

Results

In total 23 scarabaeid dung beetle species were recorded. Of the 1427 specimens sampled, the majority belonged to the genera Onthophagus and Aphodius with 714 and 710 individuals and 9 and 11 species, respectively. In addition, singletons of Oxyomus sylvestris, Rhyssemus germanus and Trypocopris vernalis were collected.

Effects of landscape parameters on species richness and abundance

We observed a positive effect of woodland cover and hedgerow length on species richness and abundance. Further, a strong positive effect of set-aside land on species richness (but not abundance) was detected. NSCP did not show any clear effect on neither richness nor abundance of beetle communities (Table 1, Figure 2).

Figure 2.

Figure 2.

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Linear regression showing significant (p = 0.05) relationships between species richness (estimated by Jack2) and abundance, and the habitat variables: (a) set-aside land, (b, d) woodland cover, and (c) hedgerow length, as predicted by a GLM (with normal error distribution and identity link-function).

Similarity and nestedness of species communities

Occurrence of dung beetle species in the 14 sampled landscapes was not random but proved to be highly nested (binmatnest: matrix T = 14.70, p < 0.001). The rank of landscapes in the presence-absence matrix packed into a state of maximum nestedness was closely related to woodland cover, but not to NSCP, set-aside land and hedgerow length (Appendix Figure A.1). The relation with crop cover did not remain significant after adjusting it for FDR (Table 2).

Table 2.

Results of Spearman rank correlations to test for relationships between the ranking of sampled landscapes in the presence-absence matrix packed into a state of maximum nestedness (Figure 3) and all landscape variables

Variables rs p FDR adjusted p
log (NSCP) − 0.16 0.5829 0.7286
arcsin (crop cover) 0.59 0.0265 0.0662
arcsin (set-aside land) − 0.43 0.1258 0.2097
arcsin (woodland cover) − 0.74 0.0023 0.0115
log (hedgerow length + 1) − 0.09 0.7705 0.7705
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Habitat variables printed bold remained significant after adjusted for false discovery rate (FDR).

Similarity relationships (quantified by Bray-Curtis similarities using square-rooted transformed abundances) between dung beetle communities of the 14 landscapes were visualized in a 2-dimensional NMDS plot (Figure 3). The plot as well as correlations between Dimension 1 values and landscapes variables indicate that changes in species composition are related to woodland cover (rs = 0.35, p = 0.025) and crop cover (rs = 0.37, p = 0.022). No relationships between set-aside land and MPAR and Dimension 1 values were found and none of the correlations between landscape variables and Dimension 2 values achieved a significant level (all p values > 0.1).

Discussion

In this study, most species were caught in landscapes with a high proportion of woodlands and hedgerow length. Areas with a high land-use intensity were characterized by an extremely depauperate dung beetle fauna. Such homogeneous landscapes dominated by habitats characterized by high land-use intensity also did not contribute to the overall species richness. Their species communities proved to be mainly subsets of landscapes with a higher proportion of woodlands and set-aside lands.

Differences of dung beetle communities between sampled landscapes may be particularly shaped by the three important factors (1) food availability, (2) habitat microclimate, and (3) soil structure (Silva and Hernández 2014). All three factors may be affected by changes of landscape composition predominantly related to land-use intensity, as intensive land use often reduces the diversity and abundance of dung-producing animals, which are essential for dung beetle sustenance (Frank et al. 2017). Additionally, intensive land use can modify habitat microclimate and soil structure by changing vegetation cover and soil compaction (Kuntzman and Brom 2024), respectively, thereby impacting the microhabitats that dung beetles rely on for breeding and feeding (Barretto et al. 2020). Most dung beetles are considered opportunistic and use a wide variety of excrement types without much discrimination (Tonelli et al. 2021). Furthermore, many species use both dung and carrion (Gimenez et al. 2021). The most important dung producing mammals in our study area are wild boar (Sus scrofa) and roe deer (Capreolus capreolus) (own observations). Both use woodland and set-aside land for foraging, resting and hiding. Therefore, landscapes with lower woodland and set-aside land are characterized by lower densities of these large mammals (Thurfjell et al. 2009; Schwegmann et al. 2023). Together with other herbivorous and carnivorous vertebrates, their dung represents the most important natural food source for dung beetles in agriculturally used areas. Empirical evidence indicates that a large and diverse mammal fauna is crucial for the maintenance of a large and diverse dung beetle fauna (Nichols et al. 2008). Consequently, a high abundance and diversity of dung producers should positively affect the richness of dung beetle communities. Dung beetles partition their food and breeding sources according to their physico-chemical attributes. These include water content, fiber size, dropping size, and nutritional quality (Tshikae et al. 2008). While the latter three parameters do exclusively depend on the type of dung producer, water content depends predominantly on the microclimate which can change dramatically between intensively used open areas (such as farmland), and particularly woodland (Nervo et al. 2021). Dung droppings in woodland and set-aside land need more time to dry out. Hence, they are longer usable and detectable for most dung beetles than droppings exposed in agricultural areas, because they are better shaded and are sheltered from wind and sun by trees and a high grass/herb layer, respectively. Therefore, the microclimate of set-aside land and woodland should have a positive effect on faunal richness of dung beetles (Righi et al. 2018; Davis et al. 2013).

Additionally, soil type and structure can have prominent effects on the occurrence of dung beetle species. Both can be important for tunnel structure and depth, and consequently for the breeding success (Bertone et. al. 2006). Unlike agriculturally used soils, forest and fallow soils offer better aeration and more structures for the protection against desiccation. Likewise, in crop areas, intensive rain is a bigger threat to the breeding chambers (Sun et al. 2023). The reason why less species are present in areas that are intensively agriculturally used could be the recurrent disturbance of the soil because of tillage (Muoni et al. 2019).

Furthermore, the widespread use of insecticides, herbicides, fungicides, and anthelmintics can be responsible for a decreasing dung beetle population (Torabian et al. 2024). Therefore, areas such as set-aside land and woodland, which are not directly affected by the applications of such chemicals, may represent refuges maintaining higher local species richness. From these “habitat islands” dung beetle species may be also able to re-colonize the adjacent farmland. In this study, increasing diversity and spatial heterogeneity of habitats quantified by NSCP and MPAR did not show any impact on species richness. Dung and carrion are ephemeral resources with a patchy distribution of space and time (Butterworth et al. 2023). Although vision is essential for navigation when flying in search for fresh dung, dung beetles primarily locate their food and breeding resources by olfactory cues that are provided by the many chemical compounds that are released from dung as volatiles, either in small or large quantities (Tshikae et. al. 2008). Dung beetles can detect food and breeding resources from more than at least 100 m. They are mobile and able to cross longer distances between different dung pats (Huerta et al. 2010). Hence, the structure and complexity of a landscape might be less vital for them than for other less mobile insects.

Conclusions

This study underscores the importance of woodland, hedgerows and set-aside land, in maintaining dung beetle diversity within agricultural landscapes. These habitats provide essential resources and favorable conditions that support diverse dung beetle communities. Further, preserving and integrating these habitats into agricultural areas is crucial for enhancing biodiversity and the ecosystem services provided by dung beetles, such as nutrient cycling and soil health. Conservation strategies should prioritize the protection and restoration of these habitats to mitigate the adverse effects of agricultural intensification. By maintaining landscape heterogeneity, it is possible to create a more resilient agricultural ecosystem that supports biodiversity and agricultural productivity. Hence, the study highlights the need for a balanced approach to land management that considers both agricultural needs and biodiversity conservation.

Acknowledgements

We would like to thank Dietmar Moser who assisted with the GIS analysis, Claudia Huber helped with the identification of dung beetles. We are grateful to all the farmers who gave their permission to work on their land. The work was financially supported by the Austrian Science Fund (Grant No. P16972).

Appendix

See Table 3, Fig. 4

Table 3.

Measured landscape variables

Landscape MPAR Crop cover (%) Set-aside land (%) Woodland cover (%)
1 0.138 39.60 9.70 33.10
2 0.246 55.10 19.70 4.20
3 0.169 61.90 11.70 1.40
4 0.240 64.10 8.50 0.50
5 0.133 67.90 1.70 0.10
6 0.068 59.30 13.10 7.40
7 0.170 49.10 29.10 2.70
8 0.044 12.00 11.30 1.90
9 0.122 41.20 38.50 14.00
10 0.183 47.50 17.30 21.40
11 0.103 64.10 8.20 2.20
12 0.171 44.20 23.90 10.20
13 0.088 60.40 14.60 0.20
14 0.056 62.90 7.40 0.30
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MPAR Mean perimeter area ratio of all land cover patches, NSCP Number of shape characterizing points

Figure 4.

Figure 4.

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Presence-absence matrices of dung beetles recorded at 14 study sites packed into a state of maximum nestedness. Study sites are in columns, species are in rows. Grey cells indicate occurrences of individual species at the sampled sites. Landscape number correspond to the numbers provided in Figure 1.

Author contributions

TF, CHS conceived the study design; CHS designed the methodology; BK did the field work; RIH, TF managed the writing of the manuscript. All authors contributed critically to the drafts and gave final approval for publication.

Funding

Open access funding provided by University of Natural Resources and Life Sciences Vienna (BOKU). This article was funded by Austrian Science Fund, P16972, P16972, P16972, P16972.

Data availability

The datasets collected and analyzed during the current study are available from the corresponding author on reasonable request.

Declarations

Competing interests

The authors declare no competing interests.

Footnotes

Publisher's Note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

References

  1. Albrecht M, Schmid B, Hautier Y, Müller CB (2012) Diverse pollinator communities enhance plant reproductive success. Proc Biol Sci 279:4845–4852 [DOI] [PMC free article] [PubMed] [Google Scholar]
  2. Babu V (2018) Effect of restoration and surrounding grassland proportion on the dung beetle metapopulation abundance in the fragmented semi-natural grasslands. Master’s thesis, Swedish University of Agricultural Sciences, Department of Ecology
  3. Barretto J, Salomão RP, Iannuzzi L (2020) Diversity of dung beetles in three vegetation physiognomies of the Caatinga dry forest. Int J Trop Insect Sci 40:385–392 [Google Scholar]
  4. Bertone MA, Green JT, Washburn SP, Poore MH, Watson DW (2006) The contribution of tunneling dung beetles to pasture soil nutrition. Forage Grazinglands 4:1–12 [Google Scholar]
  5. Blaauw BR, Isaacs R (2014) Flower plantings increase wild bee abundance and the pollination services provided to a pollination-dependent crop. J Appl Ecol 51:890–898 [Google Scholar]
  6. Bunalski M (1999) Die Blatthornkäfer Mitteleuropas (Coleoptera, Scarabaeoidea). Bestimmung − Verbreitung − Ökologie. F. Slamka, Bratislava [Google Scholar]
  7. Bürgler M, Hussain RI, Maas B, Walcher R, Rabl D, Krautzer B, Moser D, Frank T (2024) New grasslands promote pollination but not biological pest control in nearby arable fields in the short term. Arthropod Plant Interact 18:327–338 [DOI] [PMC free article] [PubMed] [Google Scholar]
  8. Butterworth NJ, Benbow ME, Barton PS (2023) The ephemeral resource patch concept. Biol Rev 98:697–726 [DOI] [PubMed] [Google Scholar]
  9. Clarke KR (1993) Non-parametric multivariate analyses of changes in community structure. Aust J Ecol 18:117–143 [Google Scholar]
  10. Clarke KR, Warwick RM (2001) Change in marine communities: An approach to statistical analysis and interpretation. Primer-E, Plymouth [Google Scholar]
  11. Davis AL, van Aarde RJ, Scholtz CH, Guldemond RA, Fourie J, Deschodt CM (2013) Is microclimate-driven turnover of dung beetle assemblage structure in regenerating coastal vegetation a precursor to re-establishment of a forest fauna? J Insect Conserv 17:565–576 [Google Scholar]
  12. Drapela T, Moser D, Zaller JG, Frank T (2008) Spider assemblages in winter oilseed rape affected by landscape and site factors. Ecography 31:254–262 [Google Scholar]
  13. Duflot R, Ernoult A, Aviron S, Fahrig L, Burel F (2017) Relative effects of landscape composition and configuration on multi-habitat gamma diversity in agricultural landscapes. Agric Ecosyst Environ 241:62–69 [Google Scholar]
  14. Escudero A, Veirano GS, González-Vainer P (2025) The applicability of a passive hole-board olfactometer in the discrimination of the presence of drugs in fecal matter for dung beetles. Chemoecology. 10.1007/s00049-025-00418-9 [Google Scholar]
  15. Fahrig L, Baudry J, Brotons L, Burel FG, Crist TO, Fuller RJ, Sirami C, Siriwardena GM, Martin JL (2011) Functional landscape heterogeneity and animal biodiversity in agricultural landscapes. Ecol Lett 14:101–112 [DOI] [PubMed] [Google Scholar]
  16. Frank K, Hülsmann M, Assmann T, Schmitt T, Blüthgen N (2017) Land use affects dung beetle communities and their ecosystem service in forests and grasslands. Agric Ecosyst Environ 243:114–122 [Google Scholar]
  17. Gardner TA, Hernández MI, Barlow J, Peres CA (2008) Understanding the biodiversity consequences of habitat change: the value of secondary and plantation forests for neotropical dung beetles. J Appl Ecol 45:883–893 [Google Scholar]
  18. Geiger F, Bengtsson J, Berendse F, Weisser WW, Emmerson M, Morales MB, Ceryngier P, Liira J, Tscharntke T, Winqvist C, Eggers S (2010) Persistent negative effects of pesticides on biodiversity and biological control potential on European farmland. Basic Appl Ecol 11:97–105 [Google Scholar]
  19. Gimenez GVC, Verdú JR, Velazco SJ, Zurita GA (2021) Dung beetle trophic ecology: are we misunderstanding resources attraction? Ecol Entomol 46:552–561 [Google Scholar]
  20. Haaland C, Naisbit RE, Bersier LF (2011) Sown wildflower strips for insect conservation: a review. Insect Conserv Divers 4:60–80 [Google Scholar]
  21. Harvey CA, Gonzalez J, Somarriba E (2006) Dung beetle and terrestrial mammal diversity in forests, indigenous agroforestry systems and plantain monocultures in Talamanca, Costa Rica. Biodivers Conserv 15:555–585 [Google Scholar]
  22. Huerta C, Anduaga S, López-Portillo J, Halffter G (2010) Use of food and space by tunneler dung beetles (Coleoptera; Scarabaeinae) during reproduction. Environ Entomol 39:1165–1169 [DOI] [PubMed] [Google Scholar]
  23. Hussain RI, Brandl M, Maas B, Rabl D, Walcher R, Krautzer B, Entling MH, Moser D, Frank T (2021) Re-established grasslands on farmland promote pollinators more than predators. Agric Ecosyst Environ 319:107543 [Google Scholar]
  24. Hussain RI, Walcher R, Vogel N, Krautzer B, Rasran L, Frank T (2023) Effectiveness of flowers strips on insect’s restoration in intensive grassland. Agric Ecosyst Environ 348:108436 [Google Scholar]
  25. Jeanneret P, Schüpbach B, Pfiffner L, Walter T (2003) Arthropod reaction to landscape and habitat features in agricultural landscapes. Landsc Ecol 18:253–263 [Google Scholar]
  26. Kessler M, Abrahamczyk S, Bos M, Buchori D, Putra DD, Robbert Gradstein S, Höhn P, Kluge J, Orend F, Saleh PR, S (2011) Cost-effectiveness of plant and animal biodiversity indicators in tropical forest and agroforest habitats. J Appl Ecol 48:330–339 [Google Scholar]
  27. Klausnitzer B (1994a) Die Käfer Mitteleuropas, Bd. 2: Myxophaga, Polyphaga 1. Spektrum Akademischer Verlag, Heidelberg [Google Scholar]
  28. Klausnitzer B (1994b) Die Käfer Mitteleuropas, Bd. 3: Polyphaga 2. Spektrum Akademischer Verlag, Heidelberg [Google Scholar]
  29. Kuntzman J, Brom J (2024) From fields to microclimate: assessing the influence of agricultural landscape structure on vegetation cover and local climate in Central Europe. Remote Sens 17:6 [Google Scholar]
  30. Landis DA (2017) Designing agricultural landscapes for biodiversity-based ecosystem services. Basic Appl Ecol 18:1–12 [Google Scholar]
  31. Martin EA, Dainese M, Clough Y, Báldi A, Bommarco R, Gagic V, Garratt MP, Holzschuh A, Kleijn D, Kovács-Hostyánszki A, Marini L (2019) The interplay of landscape composition and configuration: new pathways to manage functional biodiversity and agroecosystem services across Europe. Ecol Lett 22:1083–1094 [DOI] [PubMed] [Google Scholar]
  32. Moser D, Zechmeister HG, Plutzar C, Sauberer N, Wrbka T, Grabherr G (2002) Landscape patch shape complexity as an effective measure for plant species richness in rural landscapes. Landsc Ecol 17:657–669 [Google Scholar]
  33. Muoni T, Mhlanga B, Forkman J, Sitali M, Thierfelder C (2019) Tillage and crop rotations enhance populations of earthworms, termites, dung beetles and centipedes: evidence from a long-term trial in Zambia. J Agric Sci 157:504–514 [Google Scholar]
  34. Nervo B, Roggero A, Chamberlain D, Caprio E, Rolando A, Palestrini C (2021) Physiological, morphological and ecological traits drive desiccation resistance in north temperate dung beetles. BMC Zoo 6:1–13 [Google Scholar]
  35. Nichols E, Larsen T, Spector S, Davis AL, Escobar F, Favila M, Vulinec K, Network TSR (2007) Global dung beetle response to tropical forest modification and fragmentation: a quantitative literature review and meta-analysis. Biol Conserv 137:1–19 [Google Scholar]
  36. Nichols E, Spector S, Louzada J, Larsen T, Amezquita S, Favila ME, Network TSR (2008) Ecological functions and ecosystem services provided by Scarabaeinae dung beetles. Biol Conserv 141:1461–1474 [Google Scholar]
  37. Pike N (2011) Using false discovery rates for multiple comparisons in ecology and evolution. Methods Ecol Evol 2:278–282 [Google Scholar]
  38. Priyadarshana TS, Martin EA, Sirami C, Woodcock BA, Goodale E, Martínez-Núñez C, Lee MB, Pagani-Núñez E, Raderschall CA, Brotons L, Rege A (2024) Crop and landscape heterogeneity increase biodiversity in agricultural landscapes: a global review and meta-analysis. Ecol Lett 27:14412 [Google Scholar]
  39. Querner P, Bruckner A, Drapela T, Moser D, Zaller J, Frank T (2013) Landscape and site effects on Collembola diversity and abundance in winter oilseed rape fields in eastern Austria. Agric Ecosyst Environ 164:145–154 [Google Scholar]
  40. Righi CA, Sandoval Rodríguez C, Ferreira EN, Godoy WA, Cognato AI (2018) Microclimatic conditions for dung beetle (Coleoptera: Scarabaeidae) occurrence: land use system as a determining factor. Environ Entomol 47:1420–1430 [DOI] [PubMed] [Google Scholar]
  41. Rischen T, Kaffenberger M, Plath E, Wolff J, Fischer K (2023) Configurational landscape heterogeneity: crop-fallow boundaries enhance the taxonomic diversity of carabid beetles and spiders. Agric Ecosyst Environ 341:108194 [Google Scholar]
  42. Rodríguez-Gironés MA, Santamaría L (2006) A new algorithm to calculate the nestedness temperature of presence–absence matrices. J Biogeogr 33:924–935 [Google Scholar]
  43. Römbke J, Hempel H, Scheffczyk A, Schallnass HJ, Alvinerie M, Lumaret JP (2007) Environmental risk assessment of veterinary pharmaceuticals: development of a standard laboratory test with the dung beetle Aphodius constans. Chemosphere 70:57–64 [DOI] [PubMed] [Google Scholar]
  44. Saleh S, Schulze C, Tscharntke T (2005) Changes of dung beetle communities from rainforests towards agroforestry systems and annual cultures in Sulawesi (Indonesia). Biodivers Conserv 14:863–877 [Google Scholar]
  45. Schernhammer T (2020) Die Dungkäferfauna (Scarabaeidae) des Steinfelds–ein Best Practice-Modell für eine Dauerweide. Biodiversität und Naturschutz in Ostösterreich–BCBEA 5:17–24 [Google Scholar]
  46. Schwegmann S, Hendel AL, Frey J, Bhardwaj M, Storch I (2023) Forage, forest structure or landscape: what drives roe deer habitat use in a fragmented multiple-use forest ecosystem? For Ecol Manage 532:120830 [Google Scholar]
  47. Shah MI, Abbas S, Olohunlana AO, Sinha A (2023) The impacts of land use change on biodiversity and ecosystem services: an empirical investigation from highly fragile countries. Sustain Dev 31:1384–1400 [Google Scholar]
  48. Shi X, Ma C, de Kraker J, Gong S, Hodgson JA, Luo S, van der Steen JJ, Xiao H, Wang F, Tie X, Chen Z (2024) Influence of agricultural intensification on pollinator pesticide exposure, food acquisition and diversity. J Appl Ecol 61:1905–1917 [Google Scholar]
  49. Shi XMAC, de Kraker J, Gong S, Hodgson JA, Luo S, van der Steen JJ, Xiao H, Wang F, Tie X, Chen Z (2024) Influence of agricultural intensification on pollinator pesticide exposure, food acquisition and diversity. J Appl Ecol 61:1905–1917 [Google Scholar]
  50. Silva PGD, Hernández MIM (2014) Local and regional effects on community structure of dung beetles in a mainland-island scenario. PLoS ONE 9:e111883 [DOI] [PMC free article] [PubMed] [Google Scholar]
  51. Southwood TRE (1978) Ecological methods with the particular reference to the study of insect populations, 2nd edn. Chapman & Hall, London [Google Scholar]
  52. Sun W, Tang W, Wu Y, He S, Wu X (2023) The influences of rainfall intensity and timing on the assemblage of dung beetles and the rate of dung removal in an alpine meadow. Biol 12:1496 [Google Scholar]
  53. Thurfjell H, Ball JP, Åhlén PA, Kornacher P, Dettki H, Sjöberg K (2009) Habitat use and spatial patterns of wild boar Sus scrofa (L.): agricultural fields and edges. Eur J Wildl Res 55:517–523 [Google Scholar]
  54. Tonelli M, Giménez Gómez VC, Verdú JR, Casanoves F, Zunino M (2021) Dung beetle assemblages attracted to cow and horse dung: the importance of mouthpart traits, body size, and nesting behavior in the community assembly process. Life (Basel) 11:873 [DOI] [PMC free article] [PubMed] [Google Scholar]
  55. Torabian S, Leffler AJ, Perkins L (2024) Importance of restoration of dung beetles in the maintenance of ecosystem services. Ecol Solut Evid 5:12297 [Google Scholar]
  56. Tscharntke T, Clough Y, Wanger TC, Jackson L, Motzke I, Perfecto I, Vandermeer J, Whitbread A (2012) Global food security, biodiversity conservation and the future of agricultural intensification. Biol Conserv 151:53–59 [Google Scholar]
  57. Tschumi M, Albrecht M, Bärtschi C, Collatz J, Entling MH, Jacot K (2016) Perennial, species-rich wildflower strips enhance pest control and crop yield. Agric Ecosyst Environ 220:97–103 [Google Scholar]
  58. Tshikae BP, Davis AL, Scholtz CH (2008) Trophic associations of a dung beetle assemblage (Scarabaeidae: Scarabaeinae) in a woodland savanna of Botswana. Environ Entomol 37:431–441 [DOI] [PubMed] [Google Scholar]
  59. Walther BA, Moore JL (2005) The concepts of bias, precision and accuracy, and their use in testing the performance of species richness estimators, with a literature review of estimator performance. Ecography 28:815–829 [Google Scholar]
  60. Weibull AC, Östman Ö, Granqvist Å (2003) Species richness in agroecosystems: the effect of landscape, habitat and farm management. Biodivers Conserv 12:1335–1355 [Google Scholar]
  61. Woodcock BA, Pywell RF, Roy DB, Rose RJ, Bell D (2005) Grazing management of calcareous grasslands and its implications for the conservation of beetle communities. Biol Conserv 125:193–202 [Google Scholar]

Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Data Availability Statement

The datasets collected and analyzed during the current study are available from the corresponding author on reasonable request.

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