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Agricultural Habitat Heterogenity Impact on the European Hare (Lepus europaeus) Density in Vojvodina Region (Serbia)


Agricultural Habitat Heterogenity Impact on the European Hare (Lepus europaeus) Density in Vojvodina Region (Serbia)

Igor Ponjiger1, Milutin Kovačević1*, Vladimir Marković1, Zoran Ristić1 and Vukan Lavadinović2

1University of Novi Sad, Faculty of Sciences, Department of Geography, Tourism and Hotel Management, Trg Dositeja Obradovića 3, 21000 Novi Sad, Serbia

2University of Belgrade, Faculty of Forestry, Kneza Višeslava 1, 11030 Belgrade, Serbia


Throughout Europe agriculture intensification during the last decades has dramatically changed the structure of the farmland landscape with an intensive impact on ecosystem stability. Changes in the production led to decreased habitat heterogeneity and to decline in biodiversity, including the European hare (Lepus europaeus), especially in the lowland regions. Even though European hare mostly adapted to intensive agricultural habitats, the population densities have decreased throughout Europe since the 1960s. Previous research reported field size to be one of the most important predictors of the hare population. We aimed to assess the effect of habitat heterogeneity and structure on hare habitat selection i.e. population density. Using hunting organization census data we explored the population density and habitat preference of the European hare in spring 2020 in the hunting ground, Bačka - Bački Petrovac. The average population density of European hare significantly differed between four fractions of the hunting ground - Bački Petrovac, Kulpin, Gložan, and Maglić. Density varied from 27 in Bački Petrovac to 50 individuals per 100 ha in Maglić. The population was negatively affected by mean patch size and by the area under no vegetation during the winter period. Conservation measures should focus on enhancing habitat heterogeneity by reducing field size, and fostering sowing during the late autumn period.

Article Information

Received 13 September 2022

Revised 25 November 2022

Accepted 15 December 2022

Available online 01 March 2023

(early access)

Published 17 April 2024

Authors’ Contribution

IP, MK and VM presented the concept and designed methodology, analysed and interpreted the data. IP, VM and ZR collected data. IP and MK wrote the manuscript. MK, ZR and VL revised the manuscript. IP, MK and VL edited the manuscript. All authors have read and agreed to the published version of the manuscript.

Key words

European hare, Population density, Hunting ground, Vojvodina


* Corresponding author:

0030-9923/2024/0003-1315 $ 9.00/0

Copyright 2024 by the authors. Licensee Zoological Society of Pakistan.

This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license (


Human activity has massively changed the land cover over the past decades (Goudie, 2018). Agricultural landscapes dominate in large parts of the world, and more than a third of Earth‘s ice-free surface is covered by agricultural land (Foley et al., 2011; Ramankutty et al., 2018), which is an important habitat for a wide range of biodiversity. In Europe, which is one of the most intensely used agricultural areas (Ramankutty et al., 2008), agriculture intensification during the last decades has dramatically changed the structure of the farmland landscape with an intensive impact on ecosystem stability which caused a strong reduction of animal biodiversity (Chamberlain et al., 2000; Donald et al., 2001). Agriculture intensification led to increased yields due to larger field sizes, the use of chemicals, and the improved efficiency of machinery (Smith et al., 2004). On the other hand, this intensification led to decreased habitat heterogeneity and to decline in biodiversity (Reidsma et al., 2006), including the European hare (Lepus europaeus) as one of the most widespread and most hunted game species in Europe, especially in the lowland regions. Brown hare, as a typical game of open grasslands, during the time has adapted to agroecosystems, or landscapes cultivated by human activity due to intensive agriculture development. Due to intensive susceptibility to habitat changes caused by agricultural intensification, European hare (Lepus europaeus) may be considered a good indicator of habitat quality (Edwards et al., 2000; Smith et al., 2005; Zellweger-Fischer et al., 2011). Although European hare has adapted to agriculturally influenced habitats, hare populations have decreased across Europe since the 1960s (Smith et al., 2004; Zellweger-Fischer et al., 2011; Kamieniarz et al., 2013; Petrovan et al., 2013; Lush et al., 2014; Panek, 2018; Pavliska et al., 2018). Research about the brown hare in Serbia, especially in the Vojvodina region (Northern Serbia) where the brown hare is the most common game species (population varies between 150.000 and 300.000) shows that numerous natural and human-induced conditions (management of hunting ground, climate factors, winter, and reproduction period loses, as well as red fox abundance on the population) impact on brown hare habitat, and directly on the number of game (Panek et al., 2006; Ristić et al., 2012; Beuković et al., 2013; Popović et al., 2014; Marković et al., 2017; Kovačević et al., 2019; Ponjiger et al., 2019; Ristić et al., 2021).

So, for sustainable management of natural areas such as hunting grounds, it is necessary to analyze these conditions to conclude why the number and density of game which is one of the most important elements of a population, varies from year to year and among hunting grounds. Land cover patches analysis through anthropogenic pressures is possible to apply for many ecological and geographical applications (Marković et al., 2014). As agricultural field size is one of the most important predictors of hare habitat preference (Mayer et al., 2018; Ullmann et al., 2018) we studied the effect of habitat heterogeneity and structure on hare habitat selection i.e. population density.

Materials and Methods

In Northern Serbia the Autonomous Province of Vojvodina there is 21526.95 km² of areas considered as hunting grounds. Hunting associations manage 120 hunting grounds with an area of 20069.11 km² (Ristić et al., 2020). Our study areas were four neighboring parts of the hunting ground Bačka (Bački Petrovac (BP), Gložan (G), Kulpin (K), and Maglić (M)) located in Vojvodina province, municipality of Bački Petrovac (Fig. 1). Ministry of Agriculture, Trade, Forestry, and Water Management permits hunting organizations to manage these hunting grounds which are unfenced type. This means that the influence of the hunters consists of improving the environmental conditions of the habitat. This hunting ground is typical lowland. All four study areas mostly consisted of arable fields (BP: 88.6%, G: 98%, K: 96.6%, and M: 98.7%), tilled with maize, soy, wheat, sugar beet, potato, barley, and to a lesser degree other crops like rapeseed, hops, and tobacco. The rest of the areas consist of forest areas, orchards, pastures and meadows, water, and built-up areas. Also, notable is the very low forested areas (0.60%) and water (2.19%). The rest of the area is covered mostly by settlements and infertile land.

All hunting associations in the Vojvodina region, annually until 31st March, report the condition and number of game, including European hare, to the Ministry of Agriculture, Trade, Forestry, and Water Management and Hunting Association of Vojvodina. For this study, data were obtained from the Hunting Association of Vojvodina. In Serbia and the Western Balkan region, the census of small game such as European hare is done by directly counting individuals in 10% of the total area of each hunting ground. Counting is carried out during one day in three parts with different densities (high, medium, and low) estimated by experts and professional service of the hunting ground (Table I). After defining three equal areas of the hunting ground (which represent 10% of the total area of the hunting ground), counters which are usually hunters, are placed in a line along the counting area with the distance between each other’s 50 m. When the counters start moving to the central point, they count hares only from the defined side (usually the left side) by driving them out from the counting area. This methodology requires highly skilled organizations and a high number of hunters and it is the practice in Serbia for decades.


For each patch, we determined the land use using satellite images. All data collected for this study was imported to ArcGIS 10.1 software by ESRI to produce maps for more visual description and geo-statistical analysis. Maps were made in WGS_1984_UTM_Zone_34N Projected Coordinate System, D_WGS_1984 Datum, with Prime Meridian Greenwich. With the same software, we determined the average dimension of fields. The satellite images were taken in March, about the same time when the hare counts were done.


Table I. Results of correlation analysis.

Areas in hunting ground

Hare per 100 ha

Area size in ha

(BP) Bački Petrovac - 1



(BP) Bački Petrovac - 2



(BP) Bački Petrovac - 3



(G) Gložan - 1



(G) Gložan - 2



(G) Gložan - 3



(K) Kulpin -1



(K) Kulpin -2



(K) Kulpin -3



(M) Maglić - 1



(M Maglić - 2



(M) Maglić - 3




Statistical analysis. All analyses were performed using the IBM SPSS (ver. 23). Pearson’s correlation coefficient r and linear regression models were applied to assess the relationship between the habitat variables and brown hare abundance. To generate predictive equations for the significance, non-significant variables were removed from the final regression model.

Results and Discussion

Average hare densities vary throughout Europe. For instance in Austria, in several agricultural habitats, there were about 95 individuals per 100 ha. Contrary in the Czech Republic, a population density of 15.4 individuals per 100 ha was recorded in predominantly homogeneous habitats (Šálek, 2014). A similar result was obtained by Santilli and Galardi (2016), whose research on the structure of the habitat (variety of crops) was the most important factor that positively influenced the number of hare.

In our study area in 4 sample plots, the density of brown hare during spring 2020 was much higher than the recorded average for AP Vojvodina. According to Ristić et al. (2020) during the period 1967-2016, the density of the brown hare in the Vojvodina region was 13.15 individuals per 100 ha. The results of the spring counts showed notable differences between the four parts of the hunting ground. The number of brown hare per 100 ha in Bački Petrovac was 27, Gložan- 30.37, Kulpin- 31, and Maglić- 50.47 per 100 ha. The distinct difference between the four parts needs to be addressed knowing the fact that they are managed by the same organization applying the same game management methods and having similar habitat conditions and predator populations.

To determine the cause of this we determined the land use for each of the hunting ground parts. The results showed significant differences in the average field sizes (patch size) (it is noticeable in Fig. 1 in the North-West part of the study area) and the areas with crops in the winter period between Maglić and the rest of the hunting ground. Several studies have shown that there are negative effects on hare population in cases of large plots and smaller diversity of crops (Schröpfer and Nyenhuis, 1982; Tapper and Barnes, 1986; Lewandowski and Nowakowski, 1993; Panek and Kamieniarz, 1999; Vaughan et al., 2003; Jennings et al., 2006). Reichlin et al. (2006) found that habitat diversity is crucial, especially during summer when cereals are harvested because this is the period when hare change their diet in favor of weeds and legumes. However, this study shows this is disputable.

A strong negative correlation (r=-.901**) was found between the percentage of fields without winter crops and hare numbers (Table I). This means the hare is less abundant in areas with fewer winter crops. Similar results were already noted in previous research. Schmidt et al. (2004) found a significant positive correlation between root crops and hare abundance and a negative relationship between winter cereals. Pepin (1989) points out that the survival rate is higher for leverets born in areas where a higher percentage of winter cereals and alfalfa are grown. Genghini and Capizzi (2005) also points to the significance of winter cereals on habitat. Reichlin et al. (2006) state that winter cereals are a significant food source during the winter months, and therefore a significant factor in habitat conditions. All this attributes to the importance of winter crops for brown hare in intensive agriculture habitats.

Also, a positive correlation (r=.693*) between field size and brown hare abundance was notable. Large fields also proved to be a negative factor for the population. The main reason for this is mainly because such crops reach maturity at the same time and after harvest, the main source of food for hare is lost over a very large area (Frylestam, 1986; McLaren et al., 1997).

Contrary, all other variables shrubs, forest areas, settlements, water, orchards, and grassland were all marked as not significant. This may be because the study area is mostly agricultural land and the percentage of these areas is very small.


Table II. Results of correlation analysis.


Correlations (r)


Correlations (r)-Maglić excluded


No winter crops




not significant



not significant


not significant



not significant


not significant



not significant


not significant



not significant


not significant



not significant


not significant



not significant


not significant

Mean field size






Since it is clear there is a significant difference in hare population in Maglić and the rest of the hunting ground we tried to exclude it from the analysis to examine the strength of the relationship in other parts. Results have shown a strong negative relationship (*) between field size and hare abundance. This means that hares are more abundant in areas with smaller fields which matches some of the previous research (Schröpfer and Nyenhuis, 1982; Tapper and Barnes, 1986; Lewandowski and Nowakowski, 1993; Panek and Kamieniarz, 1999; Vaughan et al., 2003; Jennings et al., 2006). Correlation with the percentage of fields without winter crops was negative medium strong however not significant (Table II).

These results prove some of the previous studies however the case of Maglić proves that the presence of winter crops, or lack of aforementioned, significantly affects the hare population. This corresponds with the study by Smith et al. (2005) who also found that winter wheat is positively associated with hare abundance. This is because hares choose cereals when they lack other food (Tapper and Barnes, 1986; Smith et al., 2004).


Table III. Results of the multiple regression analysis.

All parts (predictor: No winter crops)

All parts (predictor: Mean field size)

Maglić excluded (predictor: Mean field size)

R square




Adjusted R square













In maglić, we found that 43.3% of the area had winter crops. This is significantly higher than the rest of the studied area (Bački Petrovac– 11.78%, Gložan– 10.93%, Kulpin– 12.86%).

This study did not determine the structure of winter crops, however, it can be noted that in the Maglić area there is a higher percentage of root crops, while in other areas winter crops are mostly wheat which corresponds with the results of the study by Schmidt et al. (2004).

To determine exactly to what extent these variables influence the number of brown hares a linear regression analysis was carried out. The results of the regression analysis also confirmed the correlation analysis where the field size was the main predictor (Table III).


Arable fields dominate agricultural land in many European countries, thereby forming the main habitat of hares. The brown hare population in Serbia is declining over the last few decades. This fact imposes the necessity for monitoring the population and finding the causes. This paper shows that the mean patch size of fields and vegetation presence during the winter period are important factors to describe habitat selection in agricultural landscapes. Hares avoid large parcels, especially without vegetation, because it does not provide high-quality forage and restricts their spatial movements. To conclude, conservation measures focusing on enhancing habitat heterogeneity by reducing field size, and fostering sowing during the late autumn period may be an effective tool to support populations of European hares in arable farmland landscapes. While this is widely known our paper also shows that there are also exceptions to these rules. Maglić hunting area is an unusual example where hare densities were significantly higher in conditions of large agricultural parcels. This can be attributed to the agricultural practices of this area, where there is a high percentage of winter crops and an especially high percentage of root crops which seem to be suitable for the brown hare population. This research also raises further questions which should be addressed in the following research. A similar approach could be used on a larger scale such as comparing several hunting grounds in a larger area.


The study received financial support from the Ministry of Education, Science and Technological Development of the Republic of Serbia (Grant No. 451-03-68/2022-14/200125).

Statement of conflict of interest

The authors have declared no conflict of interest.


Beuković, M., Beuković, D., Popović, Z., Đorđević, N., and Đorđević, M., 2013. Impact of climatic factors to the percentage of young in the population of brown hare (Lepus europaeus P.) in the Bačka district. Acta Vet., 63: 111-122.

Chamberlain, D.E., Fuller, R.J., Bunce, R.G.H., Duckworth, J.C., and Shrubb, M., 2000. Changes in the abundance of farmland birds in relation to the timing of agricultural intensification in England and Wales. J. appl. Ecol., 37: 771-788.

Donald, P.F., Green, R.E., and Heath, M.F., 2001. Agricultural intensification and the collapse of Europe’s farmland bird populations. Proc. R. Soc. Lond., B, 268: 25-29.

Edwards, P.J., Fletcher, M.R., and Berny, P., 2000. Review of the factors affecting the decline of the European brown hare, (Lepus europaeus Pallas, 1778) and the use of wildlife incident data to evaluate the significance of paraquat. Agric. Ecosyst. Environ., 79: 95-103.

Foley, J.A., Ramankutty, N., Brauman, K.A., Cassidy, E.S., Gerber, J.S., Johnston, M., and West, P.C., 2011. Solutions for a cultivated planet. Nature, 478: 337-342.

Frylestam, B., 1986. Agricultural land use effects on the winter diet of brown hares (Lepus europaeus Pallas) in southern Sweden. Mammal. Rev., 16: 157-161.

Genghini, M., and Capizzi, D., 2005. Habitat improvement and effects on brown hare (Lepus europaeus) and roe deer (Capreolus capreolus): A case study in northern Italy. Wildl. Biol., 11: 319-329.[319:HIAEOB]2.0.CO;2

Goudie, AS., 2018. Human impact on the natural environment. USA: Wiley-Blackwell.

Jennings, N., Smith, R.K., Hackländer K., Harris, S., and White, P.C.L. 2006. Variation in demography, condition and dietary quality of hares Lepus europaeus from high-density and low-density populations. Wildl. Biol., 12: 179-189.[179:VIDCAD]2.0.CO;2

Kamieniarz, R., Voigt, U., Panek, M., Strauss, E., and Niewegłowski, H., 2013. The effect of landscape structure on the distribution of brown hare Lepus europaeus in farmlands of Germany and Poland. Acta Theriol., 58: 39-46.

Kovačević, M., Marković, V., Ponjiger, I., Ristić, Z., Matejević, M., Stojsavljević, R., and Stamenković, I. 2019. Evaluation of vegetation as a habitat factor in hunting ground. Rocznik Ochrona Środowiska, 21: 85-97.

Lewandowski, K., and Nowarowski, J.J., 1993. Spatial distribution of brown hare Lepus europaeus in habitats of various types of agriculture. Acta Ther., 38: 435-442.

Lush, L., Ward, A.I., and Wheeler, P., 2014. Opposing effects of agricultural intensification on two ecologically similar species. Agric. Ecosyst. Environ., 192: 61-66.

Marković, V., Klaučo, M., Stankov, U., Jovanović, T., and Ristić, Z., 2014. Evaluation of human impact on the land cover trough landscape metrics: Nature park Šragan-Mokra Gora (Serbia). Rocznik Ochrona Środowiska, 16: 52-73.

Marković, V., Vasliljević, D.J., Jovanović, T., Lukić, T., Vujičić, M., Kovačević, M., Ristić, Z.A., Marković, S.B., Ristanović, B., and Sakulski, D., 2017. The effect of natural and human-induced habitat conditions on number of roe deer: case study of Vojvodina, Serbia. Acta Geogr. Slovenica, 57: 57-69.

Mayer, M., Ullmann, W., Sunde, P., Fischer, C., and Blaum, N., 2018. Habitat selection by the European hare in arable landscapes: The importance of small-scale habitat structure for conservation. Ecol. Evol., 8: 11619-11633.

McLaren, G.W., Hutchings, M.R., and Harris, S., 1997. Why are brown hares (Lepus europaeus) rare in pastural landscapes in Great Britain? Gibier Faune Sauvage, 14: 335–348.

Panek, M. and Kamieniarz, R., 1999. Relationships between density of brown hare Lepus europaeus and landscape structure in Poland in the years 1981-95. Acta Theriolog., 44: 67-75.

Panek, M., 2018. Habitat factors associated with the decline in brown hare abundance in Poland in the beginning of the 21st century. Ecol. Indic., 85: 915-920.

Panek, M., Kamieniarz, R., and Bresinski, W., 2006. The effect of experimental removal of red foxes Vulpes vulpes on spring density of brown hares Lepus europaeus in western Poland. Acta Theriol., 51: 187-193.

Pavliska, P.L., Riegert, J., Grill, S., and Salek, M., 2018. The effect of landscape heterogeneity on population density and habitat preferences of the European hare (Lepus europaeus) in contrasting farmlands. Mammal. Biol., 88: 8-15.

Pepin, D., 1989. Variation in survival of brown hare (Lepus europaeus) leverets from different farmland areas in the Paris basin. J. appl. Ecol., 26: 13-23.

Petrovan, S.O., Ward, A.I., and Wheeler, P.M., 2013. Habitat selection guiding agri-environment schemes for a farmland specialist, the brown hare. Anim. Conserv., 16: 344-352.

Ponjiger, I., Ristić, Z., Marković, V., Matejević, M., and Kovačević, M., 2019. The dynamics of red fox (Vulpes vulpes) and brown hare (Lepus europaeus) populations in the Vojvodina region (Serbia) in relation to rabies vaccination. Vet. Arh., 89: 839-850.

Popović, Z., Beuković, M., and Beuković, D., 2014. Management measures in brown hare population in various habitats in Serbia. In: Proceedings of the international symposium on animal science (ed. Z. Popović). Faculty of Agriculture, Belgrade. pp. 428-437.

Ramankutty, N., Evan, A.T., Monfreda, C., and Foley, J.A., 2008. Farming the planet: 1. Geographic distribution of global agricultural lands in the year 2000. Glob. Biogeochem. Cyc., 22: GB1003.

Ramankutty, N., Mehrabi, Z., Waha, K., Jarvis, L., Kremen, C., Herrero, M., Rieseberg, LH., 2018. Trends in global agricultural land use: Implications for environmental health and food security. Annls Rev. Pl. Biol., 69: 789-815.

Reichlin, T., Klansek, E., and Hackländer, K., 2006. Diet selection by hares (Lepus europaeus) in arable land and its implications for habitat management. Eur. J. Wildl. Res., 52: 109-118.

Reidsma, P., Tekelenburg, T., Van den Berg, M., and Alkemade, R., 2006. Impacts of land-use change on biodiversity: An assessment of agricultural biodiversity in the European Union. Agric. Ecosyst. Environ., 114: 86-102.

Ristić, Z., Marković, V., Barović, V., and Vasiljević, D.J., 2012. Loss calculation model of brown hare (Lepus europaeus Pall) and its application in the hunting grounds of Vojvodina region (north Serbia). Pakistan J. Zool., 44: 1-5.

Ristić, Z., Ponjiger, I., Matejević, M., Kovačević, M., Ristić, N., and Marković, V., 2021. Effects of factors associated with the decline of brown hare abundance in the Vojvodina region (Serbia). Hystrix Ital. J. Mammal., 32: 67-71.

Ristić, Z., Urošević, I.M., Ristić, N., Matejević, M., Ponjiger, I., Kovačević, M., and Ćirović, P., 2020. Planned and rational management of the brown hare (Lepus europaeus Pallas, 1778) population in Vojvodina region (Serbia) for the past 50 years (1967-2016). Balkan J. Wildl. Res., 5: 28-35.

Santilli, F., and Galardi, L., 2016. Effect of habitat structure and type of farming on European hare (Lepus europaeus) abundance. Hystrix Ital. J. Mammal., 27: 120-122.

Schmidt, N.M., Asferg, T., and Forchhammer, M.C., 2004. Long-term patterns in European brown hare population dynamics in Denmark: effects of agriculture, predation and climate. BMC Ecol., 4:

Schröpfer, R., Nyenhuis, H. 1982. Die Bedeutung der Landschaftsstruktur für die Populationsdichte des Feldhasen (Lepus europaeus Pallas, 1778). Z. Jagdwissenschaft, 28: 213-231.

Smith, R.K., Jennings N.V., and Harris, S., 2005. A quantitative analysis of the abundance and demography of European hares Lepus europaeus in relation to habitat type, intensity of agriculture and climate. Mammal. Rev., 35: 1-24.

Smith, R.K., Jennings, N.V., Robinson, A., and Harris, S., 2004. Conservation of European hares Lepus europaeus in Britain: Is increasing habitat heterogeneity in farmland the answer? J. appl. Ecol., 41: 1092-1102.

Šálek, M., 2014. Farmland biodiversity: Searching for determinants of species diversity and abundance in farmlands with contrasting habitat heterogeneity. Final Report, Institute of Vertebrate Biology AV ČR, Brno, pp. 81.

Tapper S.C., and Barnes, R.F.W., 1986. Influence of farming practice on the ecology of the brown hare (Lepus europaeus). J. appl. Ecol., 23: 39-52.

Ullmann, W., Fischer, C., Pirhofer-Walzl, K., Kramer-Schadt, S. and Blaum, N., 2018. Spatiotemporal variability in resources affects herbivore home range formation in structurally contrasting and unpredictable agricultural landscapes. Landscape Ecol., 33: 1505-1517.

Vaughan, N., Lucas, E.A., Harris, S. and White, P.C.L., 2003. Habitat associations of European hares Lepus europaeus in England and Wales: Implications for farmland management. J. appl. Ecol., 40: 163-175.

Zellweger-Fischer, J., Kéry, M. and Pasinelli, G., 2011. Population trends of brown hares in Switzerland: the role of land-use and ecological compensation areas. Biol. Consv., 144: 1364-1373.

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