Acta geographica Slovenica, 57-2, 2017, 57–69
THE EFFECT OF NATURAL
AND HUMAN-INDUCED HABITAT
CONDITIONS ON NUMBER OF ROE DEER:
CASE STUDY OF VOJVODINA, SERBIA
Vladimir N. Marković, Djordjije A. Vasiljević, Tamara Jovanović, Tin Lukić,
Miroslav D. Vujičić, Milutin Kovačević, Zoran A. Ristić, Slobodan B. Marković,
Branko Ristanović, Dušan Sakulski
Roe deer in natural habitat.
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The effect of natural and human-induced habitat conditions
on number of roe deer: case study of Vojvodina, Serbia
DOI: http://dx.doi.org/AGS.903
UDC: 913:591.522(497.113)
591.522:599.735.341(497.113)
COBISS: 1.01
ABSTRACT: Roe deer (Capreolus capreolus L.) have greatly expanded in both distribution and abundance
during the last few decades, and are the most abundant cervids in Europe today. The aim of this paper is
to determine the factors that have the most considerable impact on roe deer numbers in the Vojvodina
region (North Serbia). Environmental (area in ha, total area of forest and total area of meadows and pas-
tures in hunting ground) and anthropogenic (number of registered hunters, number of hunting sections,
number of gamekeepers and roads in km on 1,000 ha) factors had been shown to influence the number
of roe deer in Vojvodina region. A multiple regression analysis was carried out as the main statistical approach.
The mapping of certain parameters was done using ArcGIS 9.2 software in order to establish the relation
between the roe deer population and the different environmental and anthropogenic conditions. The results
signify that the roe deer number dependency in the Vojvodina region is a very complex and multi-facto-
rial phenomenon, strongly influenced by human induced modifications. 
KEY WORDS: geography, environmental protection, roe deer number, habitat, hunting, regression model,
GIS, Vojvodina, Serbia
The article was submitted for publication on August 8th, 2014.
ADRESSES:
Vladimir N. Marković, Ph.D.
Department of Geography, Tourism and Hotel Management
Faculty of Sciences, University of Novi Sad
Trg Dositeja Obradovića 3, RS – 21000 Novi Sad, Serbia
E-mail: vladimir.markovic@dgt.uns.ac.rs
Djordjije A. Vasiljević, Ph.D.
Department of Geography, Tourism and Hotel Management
Faculty of Sciences, University of Novi Sad
Trg Dositeja Obradovića 3, RS – 21000 Novi Sad, Serbia
E-mail: vasiljevic80@gmail.com
Tamara Jovanović, Ph.D.
Department of Geography, Tourism and Hotel Management
Faculty of Sciences, University of Novi Sad
Trg Dositeja Obradovića 3, RS – 21000 Novi Sad, Serbia
E-mail: tamara.jovanovic@dgt.uns.ac.rs
Tin Lukić, Ph.D.
Department of Geography, Tourism and Hotel Management
Faculty of Sciences, University of Novi Sad
Trg Dositeja Obradovića 3, RS – 21000 Novi Sad, Serbia
E-mail: tin.lukic@dgt.uns.ac.rs
The effect of natural and human-induced habitat conditions on number of roe deer: case study of Vojvodina, Serbia
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59
Acta geographica Slovenica, 57-2, 2017
Miroslav D. Vujičić, Ph.D.
Department of Geography, Tourism and Hotel Management
Faculty of Sciences, University of Novi Sad
Trg Dositeja Obradovića 3, RS – 21000 Novi Sad, Serbia
E-mail: miroslav.vujicic@dgt.uns.ac.rs
Milutin Kovačević M.Sc.
Department of Geography, Tourism and Hotel Management
Faculty of Sciences, University of Novi Sad
Trg Dositeja Obradovića 3, RS – 21000 Novi Sad, Serbia
E-mail: milutin.kovacevic@dgt.uns.ac.rs
Zoran A. Ristić, Ph.D.
Department of Geography, Tourism and Hotel Management
Faculty of Sciences, University of Novi Sad
Trg Dositeja Obradovića 3, RS – 21000 Novi Sad, Serbia
E-mail: balzakova@yahoo.com
Slobodan B. Marković, Ph.D.
Department of Geography, Tourism and Hotel Management
Faculty of Sciences, University of Novi Sad
Trg Dositeja Obradovića 3, RS – 21000 Novi Sad, Serbia
E-mail: slobodan.markovic@dgt.uns.ac.rs
Branko Ristanović, Ph.D.
Department of Geography, Tourism and Hotel Management
Faculty of Sciences, University of Novi Sad
Trg Dositeja Obradovića 3, RS – 21000 Novi Sad, Serbia
E-mail: branko.ristanovic@dgt.uns.ac.rs
Dušan Sakulski, Ph.D.
Department of Environmental Engineering and Occupational Safety and Health
Faculty of Technical Sciences, University of Novi Sad
Trg Dositeja Obradovića 6, RS – 21000 Novi Sad, Serbia
and Disaster Management Training and Education Centre (DiMTEC)
University of the Free State
205 Nelson Mandela Drive, Park West, ZA – 9300 Bloemfontein, South Africa
E-mail: dsakulski2@me.com
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Figure 1: The map of roe deer dissemination in the Vojvodina region (Antonić and Beuković 2007; modified). p
60
1 Introduction
During the last few decades, roe deer (Capreolus capreolus L.) have greatly expanded in both distribution
and abundance, and today they are the most abundant cervids in Europe (Andersen, Duncan and Linnel
1998; Mysterud and Østbye 2004; Apollonio, Andersen and Putman 2010; Torres et al. 2011) as well as
the most extensively studied ungulates (Andersen, Duncan and Linnell 1998).
Although generally considered as a typical forest specie, recent studies proved roe deer’s ability to inhabit
physically diverse areas and could be found in almost all European landscapes (Hewison et al. 2001; Jepsen
and Topping 2004, Torres et al. 2011).
The Vojvodina region (North Serbia) is characterised by vast areas of steppe, wetlands, ponds and forests,
which are favourable habi-tats for numerous and diverse game species (Ristić, Marković and Dević 2009).
The number of roe deer within the investigated area varies among hunting associations. According to the
data from 2000, the average number of roe deer in Vojvodina is 17.6 heads on 1,000 ha. The long-term
hunting development plan 2001–2010 (Hunting association … 2000), predicted a 49.03% rise in the num-
ber of roe deer, which should bring the number of roe deer to 26.2 heads on 1,000 ha in 2010. The data
from 2010 are not currently available, but according to the last available data, from 2009, the number has
raised to 25.4 heads on 1,000 ha. The aim of this paper is to determine the factors with the most consid-
erable impact on the number of roe deer in the investigated area. Although this species’ habitat is widely
influenced by a range of biogeographical factors operating at different scales (Torres et al. 2011), recent
researches have determined that available food (Virgós and Telléria 1998) and land cover (Mysterud and
Østbye 1999; Borkowski 2004; Borkowski and Ukalska 2008; Melis et al. 2009) are two essential factors.
These factors play a critical part in shaping this species’ habitat selection, which in turn affects the num-
ber of roe deer. However, Torres et al. (2011) consider several other factors that could also affect its population,
including human interference and disturbance (Aragón, Braza and San José 1995; Hewison et al. 2001),
topography (Mysterud and Østbye 1999), and climatic factors (Brewka and Kossak 1994). This research
combines quantitative data on certain factors suggested to have the biggest impact on roe deer habitat suit-
ability in order to determine the most influential ones. Environmental (area in ha, total area of forest and
total area of meadows and pastures in hunting ground) and anthropogenic (number of registered hunters,
number of hunting sections, number of gamekeepers and roads in km on 1,000 ha) factors influence the
number of roe deer in Vojvodina region.
2 Materials and methods
2.1 Method
2.1.1 Sample
The study area includes 57 hunting grounds (Figure 1) that correspond with the municipality areas – except
for the Kikinda, Vršac, Srbobran and Ada municipalities with more than one hunting ground on their area.
The hunting grounds are unfenced and managed by the Provincial Secretariat for Agriculture, Forestry
and Water Management. The game on these hunting grounds is self-reliant and only influenced by human
induced environmental improvements. The following hunting grounds were excluded from the study because
they are fenced off and supports different habitat conditions: The hunting grounds of National Park »Fruška
gora«; hunting grounds in possession of military of Serbia; Public Company »Vojvodina šume«; and hunt-
ing grounds owned by agricultural and fish ponds companies.
2.1.2 Hypotheses
The following hypotheses were postulated:
1) The number of roe deer is influenced by natural factors: Total area of hunting ground in hectares; total
area of forest on hunting ground in hectares; and total area of meadows and pastures on hunting ground
in hectares.
The effect of natural and human-induced habitat conditions on number of roe deer: case study of Vojvodina, Serbia
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All above listed natural factors positively influence the number of roe deer. Numerous researches
(e.g. Mysterud, Lian and Hermann 1999); Bokalo 2001; Partl 2001; Partl 2002; Borkowski 2004; Hemani,
Watkinson and Dolman 2005; Toïgo et al. 2008; Popović et al. 2009; Reimosera et al. 2009) support this claim.
2) The number of roe deer is influenced by human induced factors: Number of registered hunters; number
of hunting sections; number of gamekeepers; and roads in km on 1,000 ha of hunting ground.
The number of registered hunters, number of hunting sections, number of residents in the munici-
pality and roads in km on 1,000 ha of hunting ground have a negative influence on the number of roe deer.
The number of gamekeepers correlates positively with the number of roe deer. Many researchers have exam-
ined these factors and their impact on the number of game (e.g. Stedman et al. 2004; Blumstein et al. 2005;
Borkowski and Ukalska 2008; Jayakody et al. 2008; Stankowich 2008; Torres et al. 2011).
2.1.3 Procedure
This study used data on the condition and number of game (including roe deer) obtained from the labo-
ratory of Hunting Association of Vojvodina. Beside the number of game, the data also include: vegetation
structure variability; infrastructure changes; various demographic data (hunters’ number, age, gender); and
other variables that could influence the game habitat and number. In order to establish the relation between
roe deer population and different environmental and anthropogenic conditions, mapping of certain para-
meters was done using ArcGIS 9.2 software.
2.1.4 Statistical approach
A multiple regression analysis was carried out using SPSS 17 software for statistical computing. Before the
analysis, three hunting grounds were excluded because their number of roe deer was either too small or
two large in comparison with the other hunting grounds (± 2 standard deviations). A log-transformation
of the dependent variable number of roe deer was performed in order to normalise its distribution. Predictor
variables were: Total area of hunting ground in hectares; number of registered hunters; number of hunt-
ing sections; number of gamekeepers; total area of forest on hunting ground in hectares; total area of meadows
and pastures on hunting ground in hectares; and roads in km on 1,000 ha of hunting ground. The crite-
rion variable was the log-transformed number of roe deer.
3 Results
The results of the above mentioned multiple regression analysis are shown in table 1. The analysis of the
results shows that the overall regression model is significant (r = 0.648; r2 = 0.420; F(6.47) = 4.071; p < 0.01).
Only the area in ha, number of registered hunters and number of hunting sections show a statistically sig-
nificant influence. The area of the hunting ground and number of registered hunters has a significant positive
impact on the number of roe deer. However, the number of hunting sections has a significant negative impact
on the number of roe deer. No effect of a multicollinearity problem is detected in the model as all the VIF
scores are smaller than 10 and Tolerance values are larger than 0.1 (Figure 2).
4 Discussion
The results show that a larger surface area has a larger number of roe deer (Table 1). The smaller the area,
the more negatively it affects game habitat in terms of seasonality, lack of shelters, and uniformed food
supplies – all of which contribute to the decrease in the number of roe deer. According to Law on Game
and Hunting (Zakon o divljači i lovstvu 2010), a minimum of 20% of each hunting ground must be pro-
tected as reserves, where all human activities related to hunting and animal disturbance are prohibited.
This ensures safer reproducing conditions and protection of the roe deer offspring. Larger hunting ground
(HG) will evidently have larger reserves. Commonly, each of the hunting grounds in the Vojvodina region
is within only one municipality and is managed by its hunting association (Figure 3).
As there is no universal management model, each hunting ground operates independently, resulting
in temporal and spatial discordance. Hunters’ ethical behaviour is under question because there is no integral
and internal control – it varies between sections and could lead to increased poaching and illegal hunting.
This has all been proven in the regression model (Table 1).
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Table 1: The regression model of certain factors influencing the number of roe deer of the investigated area.
β (weight) t-test p (significance) Tolerance VIF
Total hunting ground area in ha 0.488 2.498 0.02 0.337 2.96
Number of registered hunters
without hunter-tourists 0.543 2.778 0.01 0.337 2.97
Number of semi-independent
hunting clubs within the hunting -0.524 -2.46 0.02 0.284 3.52
associations
Number of gamekeepers
without volunteers 0.05 0.299 0.77 0.466 2.15
roads in km on 1,000 ha
of hunting ground -0.018 -0.13 0.9 0.665 1.5
total area of meadows and pastures
on hunting ground in hectares 0.135 1.117 0.27 0.884 1.13
total area of forest on the hunting
ground in hectares -0.04 -0.3 0.77 0.712 1.4
Empirical
data from
the
laboratory
Selected
factors that
influence roe
deer habitat
Multiple
regression
analysis
Hunting
ground area
in ha
Number
of registered
hunters
Number
of hunting
sections
Number
of gamekeepers Traffic
Percentage
of green
area
Number
of roe deer
Forestation
0.488* 0.543** –0.524*
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Figure 2: Graphic presentation of regression model.
Figure 3: Quantification of hunting factors by hunting grounds (Antonić and Beuković 2007; modified). p p. 64
Figure 4: Traffic network density (km/1,000 ha) by hunting grounds in the Vojvodina region (Source: Statistical Office of the Republic of Serbia 2008;
modified) p p. 65
Figure 5: Forestation and area of meadows and pastures of hunting grounds in the Vojvodina region (Statistical Office of the Republic of Serbia 2008;
modified). p p. 66
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Roe deer generally avoid areas that are intensively used by humans (Mysterud, Lian and Hermann 1999).
However, this study found that the number of registered hunters is directly proportional to the number
of roe deer game. This might be due to increased income generated by the hunter’s membership fees that
is used for game food and medicaments, breeding and feeding facilities. Also, larger hunter numbers result
in better and more efficient care of game, especially concerning annual procedures such as game counting,
feeder and salt lick instalment, maintenance and repair of breeding and feeding facilities, or even game
rescue during natural hazards (floods, fires, snow, etc.). The role of hunters in these situations is crucial,
as hunting ground quality depends on their assistance. According to Marković et al. (2011), most of the
registered hunters in Vojvodina (46.7%) have stated that linking with nature is the primary motive for engag-
ing in hunting.
In contrast to number of hunters, number of gamekeepers does not affect the number of roe deer game.
We believe that this is mostly the case because in the investigated area most of the gamekeepers work vol-
untarily and thus their commitment to duty could be questionable. Consequently, the latest Law on Game
and Hunting (Zakon o divljači i lovstvu 2010) enacted by Ministry of Agriculture, Trade, Forestry and Water
Management of the Republic of Serbia, revoked the title gamekeeper-volunteer. According to Marković
(2010), the level of gamekeeper-volunteers’ adequate expertise and knowledge in Vojvodina has shown
considerable variability. 
The length of roads in the investigated area has no effect on roe deer number, although it was expect-
ed to have a negative influence (e.g. Mysterud 1999; Hewison et al. 2001; Torres et al. 2011). This could
be justified by the fact that average traffic network density in the Vojvodina region is only 2.93 km/1,000 ha
(of hunting ground) (Figure 4).
The total area of meadows and pastures on hunting ground in hectares, and total area of forest on hunt-
ing ground in hectares are insignificant parameters (Table 1), unlike in the findings of previous researches
(e.g. Reimoser and Zandl 1993; Partl 2001 and 2002; Bokalo 2001; Borkowski and Ukalska 2008;
Reimosera et al. 2009). Vojvodina is one of the European regions with poorest forest areas (only 5.5% of
the territory is covered by forest) (Figure 5). Furthermore, most of the forests are located within nature
reserves (outside of hunting grounds), which leaves investigated area with average forestation of only 1.7%
(Marković 2010). On the other hand, comparing three forest management models – no harvest, single tree
selection and clear-cut – the clear cut gives the best habitat quality for roe deer, which indicate that foresta-
tion is not necessarily an essential parameter for roe deer habitat (Vospernik and Reimoser 2008). We also
believe that the area of meadows and pastures is an insignificant predictor for the number of roe deer, most-
ly due to vast areas under agricultural crops with addition of feeding facilities that provide sufficient food
supply for roe deer population.
5 Conclusion
This study’s results and discussions signify that the roe deer number dependency in the Vojvodina region
is a very complex and multi-factorial phenomenon, but strongly influenced by human-induced modifica-
tions of habitat. Unfortunately, as a typical Pannonian landscape with a high level of cultivation (Antonić
and Beuković 2007), the Vojvodina region offers limited conditions for wildlife habitat (Suchant, Baritz
and Braunisch 2003). 
These agricultural regions, or »agroecosystems« (Knoche and Lupi 2004) present the most important and
pervasive type of managed ecosystem (Antle and Capalbo 2002). It is located on the most productive land, cov-
ering 30% of the earth's land area, and 80% in the Vojvodina region (Antonić and Beuković 2007). Consequently,
the establishment of agriculture and hunting grounds could influence natural habitats (Knoche and Lupi 2004),
with certain conflicts arising between wildlife management and protection on the one hand, and the utilisa-
tion of the land for tourism and economic exploitation on the other (Suchant, Baritz and Braunisch 2003).
Inevitably, there is a formal opinion that (particularly in Vojvodina region) roe deer and ungulates in
general, with habitats outside protected areas, could not survive in those numbers if there were no hunting
grounds. A great danger to the roe deer population could emerge with uncontrolled hunting, cold and dry
periods with poor land cover, food and water supplies, increased cultivated areas, etc. All of these factors
are generally prevented by the efficient and devoted work of hunting associations with the application of
adequate biogeographical surveys and strategies.
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68
ACKNOWLEDGEMENT: This work was supported by the Ministry of Science and Technological
Development, Republic of Serbia (grant 176020) and the Provincial Secretariat for Science and Technological
Development, Vojvodina Province, Serbia (grant 114-451-2644/2012-01). 
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