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1
Spatio-temporal distribution of European bison
(Bison bonasus L.) in Poloniny National Park, East
Carpathians, Slovakia
Abstract. This study focused on analysis of spa-
tio-temporal distribution of endangered European
bison in Poloniny National Park (PNP). The main
aim of the study was to analyse bison territory
during both winter and the vegetation season for
the whole period of reintroduction to show differ-
ences in occurrence at different altitudes, slopes
and aspects as well as relationships between them
during seasons and changes in territory utilization
during 3-year periods. The animals change their
habitat requirements according to actual season.
Bison visit higher altitudes and steeper slopes
mainly during the summer. During winter, bison
remain in the valleys of displaced villages and feed
on apple orchards and supplementary hay. The
population is conservative in their territory and it
still visits the same area.
Key words: altitudinal movements, habitat utilization, sea-
sonal distribution, Poloniny National Park, East Carpathians
Introduction
The European bison (Bison bonasus) is Europe’s
largest terrestrial mammal. Pucek et al. (2004) state
that in the past European bison were distributed all
over the European continent, specically in south-
eastern, central and western Europe.
In Slovakia, the last bison was killed in the 16th-
17th century (Pčola and Adamec 2007). Only two
populations of European bison survived in the wild
to the end of the 19th century; one in the Białowieża
Forest at the borders between Poland and Belarus
(B. b. bonasus) and the other in the West-Caucasus
Mountains (B. b. caucasicus) (Pucek et al. 2004).
History of the European bison extinction was sum-
marised in studies by Pucek (1991, 1994). In many
European countries the reintroduction of European
bison is accomplished using animals previously
kept in zoological enclosures (Pucek et al. 2004;
Eliáš 2006, 2008). Release of captive animals into
the wild to restore wild-living populations was
at rst attempted in the Polish part of Białowieża
Forest in 1952 (Krasiński 1978, 1983) and then in
forests in Belarus, Lithuania, Russia, Slovakia,
Ukraine (Raczyński 2011, Pucek et al. 2004) and
Kyrgyzstan (Pucek et al. 2004). However, all free-
ranging as well as semi-free herds are isolated and
these small populations quickly lose their genetic
heterogeneity and are more vulnerable to extinc-
tion (Franklin 1980). High genetic similarity in all
wild-living populations is indicative of inbreeding
as the result of the bottleneck effect. Several stud-
ies reported the high (and still rising) inbreeding co-
efcient in European bison from Białowieża Forest
(Matuszewska et al. 2004, Wołk and Krasińska 2004).
Inbreeding has a detrimental effect on health, par-
ticularly on skeleton growth in females, and reduces
disease and pathogen resistance of bison popula-
tion (Kobryńczuk 1985, Olech 2008). Therefore, it is
necessary to create conditions to facilitate migration
of bison between herds. The rst step should be to
search for natural migratory routes and space use by
individuals at sites of potential interconnections be-
tween neighbouring Slovak and Polish bison herds.
The State Nature Conservancy of the Slovak Re-
public developed and in 2004 implemented a project
entitled “The Foundation of wild-living population
of European bison in Slovak Carpathian Mountains,
Poloniny National Park” (Adamec 2004, 2006; Eliáš
2008). This project aimed to create a stable popula-
tion of European bison in Slovakia (Fig. 1). Five indi-
viduals from Italy, the Netherlands and Switzerland
were imported in June 2004 and on December 10th,
2004 were released into the wild and monitored by
telemetry (Adamec and Pčola 2004; Perzanowski et
al. 2006; Pčola and Adamec 2007; Eliáš 2006, 2008).
Shortly after their release, (December 25, 2004) one
M. BIELIK, M. JANIGA and Z. BALLOVÁ
Institute of High Mountain Biology University
of Žilina, Tatranská Javorina 7, SK-059 56, Slovakia,
e-mail: zuzana.ballova@uniza.sk
Oecologia
Montana 2018,
27, 1-6
© 2018 Prunella Publishers
Fig. 1. The European bison in the Poloniny National Park,
Slovakia (Photo: M. Janiga, 2013).
2
M. Bielik, M.
Janiga & Z.
Ballová
wild-living bull called Pišta joined the group (Ada-
mec 2006; Perzanowski et al. 2006; Pčola and Ada-
mec 2007; Eliáš 2008). The chronology of reintro-
duction in Poloniny National Park was summarised
in a study by Pčola and Gurecka (2008).
Different individuals often use the same area
and their territories overlap considerably. Central
parts of territories, including meadows and water-
ing holes are the most intensively utilised (Krasiński
et al. 1999). In Lithuania, animals frequented an
area between 100 and 200 km2 with large core areas
measuring approximately 20 km2 (Balčiauskas 1999;
Pucek et al. 2004). A satisfactory area to sustain a
population of 50-70 animals looks to be approxi-
mately 200 km2 (Pucek et al. 2004).
The territories of European bison in Slovakia
were analysed during the rst four years following
reintroduction (December 10th, 2004 to August 3rd,
2008) and mentioned in studies of Adamec (2006),
Pčola and Adamec (2007) and Pčola and Gurečka
(2008). At this time relationships were observed
between distribution of European bison herds and
individual animals and their occurrence at different
altitudes, slopes and aspects during the year, but
were never completely analysed. There are only a
few free-ranging and semi-free herds of European
bison in the world, and they occur in the wild only
in a few places (Pucek et al. 2004). Therefore, the
sustainable management of each population is key
to survival of the species on a global scale.
Research on habitat utilization and patterns of
herd movement during the whole period of popula-
tion existence are essential to improving the rein-
troduction and the long-term management of Euro-
pean bison in European reserves.
We decided to compare winter and summer
area of European bison and changes in their di-
mensions during years after reintroduction. The
main objective of this paper is to analyse spatio-
temporal distribution and identify potential con-
nection of Slovak herd with Polish herd during the
period of reintroduction of European bison popula-
tions in the Poloniny National Park, East Carpath-
ians, Slovakia since 2004 to 2017.
Material and Methods
Poloniny National Park is located in north-eastern
Slovakia, at the border with Poland and Ukraine, in
the Bukovské vrchy mountain range. Areas of PNP
(Poloniny National Park) are included in the East Car-
pathian Biosphere Reserve and several locations have
been proclaimed by UNESCO as World Heritage Sites.
Data was collected between December 10th,
2004 and February 17th, 2017 (including 13 winter
periods (December to March) and 12 vegetation
periods (April to November)). It was collected by
staff from The State Nature Conservancy of the
Slovak Republic, the Administration of PNP, lo-
cated in Stakčín and university students through
telemetric monitoring. Monitoring was conducted
once per week using the ICOM IC-R 10 receiver a
Telonics RA 14 antenna (Pčola and Gurečka 2008,
Eliáš 2008), and random observers. Between 2016
and 2017, data from authors’ regular observa-
tions (several days each month) was added. Po-
sition data of herd and solitary individuals was
collected, including date and coordinates as well
as abundance and gender when possible (mainly
since year 2010). Visual observations and eld
remnants (fresh droppings, footprints, lying ar-
eas, nibbled trees and shrubs) were recorded.
Coordinates of hay feeders in the area, used as a
source of food during winter were also recorded.
Geographic information software (ArcGIS 10.0)
was used for data analysis, specically the Ar-
cMap component. The data of basic matrix was
analysed and standardized using the STATISTI-
CA 12.1 program. Two-way analysis of variance
(ANOVA) was used to determine signicant dif-
ferences between the means of two or more inde-
pendent groups. The data is expressed as mean
± standard deviation (SD). Values of P less than
0.05 were admitted as statistically signicant.
Results
540 out of a total of 560 position coordinates were
used for analysis of area of territory. 20 coordinates
(3.6%) were not used (11 of the Polish herd, one of
migrating bull near Zboj village and 8 of migrating
bull near Stakčín village). The Polish herd was ob-
served in period between January 2016 and Febru-
ary 2017 and we found potential interconnection
between Slovak and Polish bison herds near Hostovi-
cký creek. However, direct encounters of bison from
two transboundary herds were not recorded (Fig. 2).
540 position coordinates were used for analysis
during whole period. Total area included in analysis
of the reintroduction period is 106.79 km2 (Fig. 2).
If we include the Polish herd and Slovak migrating
bulls, the total area of European bison occurrence in
the wild in Slovakia is 265.19 km2. Maximal distance
of occurrence data was 31.45 km (Svetlice – Zboj).
218 position coordinates were recorded during
the period between October and March. The total
area recorded in all winter periods was 81.33 km2.
Maximal distance of occurrence data in winter
was 12.1 km (Fig. 3a).
322 position coordinates were recorded during
the period between April and September. The total
area recorded in all vegetation periods was 104.27
km2. Maximal distance of occurrence data in veg-
etation period was 15.1 km (Fig. 3b).
In spring, eastern slopes were the most occu-
pied by bison (39.3%), followed by western (23.6%),
southern (20.2%) and northern slopes (16.9%) (Fig. 4).
In summer, western slopes were the most occupied
(28.0%), followed by southern (27.2%), eastern (26.4%)
and northern slopes (18.4%). In autumn, western
slopes were the most occupied (29.7%), followed by
eastern (26.6%), southern (25.9%) and northern slopes
(17.7%). In winter, eastern slopes were the most oc-
cupied (45.5%), followed by western (28.7%), southern
(15.7%) and northern slopes (10.1%).
There were no seasonal differences observed be-
tween altitude of occurrence of herds and solitary
animals. Herds inhabited lower altitudes in spring
and then moved to higher altitudes. Conversely, in-
dividual animals inhabited higher altitudes in spring
than herds and then moved to lower altitudes during
summer. Nevertheless, interaction between these
factors was deemed statistically insignicant.
3
Spatio-temporal
distribution of the
European bison in
the Poloniny Na-
tional Park
Fig. 2. Map of total area of the European bison occurrence in Poloniny National Park. Squares are data of Polish herd,
which occurred on Slovak territory since January 2016 and triangles are data of Slovak migrating bulls. The data was not
included to calculation of area of territory. Line encloses core area of bison occurrence in PNP.
Fig. 3. Maps of territories of European bison in Poloniny National Park in a) winter season (October to March) and b) veg-
etation season (April to September). Thick lines show winter/summer area and thin lines total area of bison occurrence.
Crosses show hay feeders. Data of Polish herd and Slovak migrating bulls are not included.
Fig. 4. Relationship between seasons, aspect and altitude. Southern and western slopes at higher altitudes from
620 to 650 m a.s.l. were commonly occupied by European bison in summer season (Two way ANOVA; effects: F(9.
534) = 3.0178; p = .00159; vertical bars denote 0.95 condence limit). In spring, autumn and winter, bisons were at
altitudes from 440 to 520 m a.s.l.
amount of food and higher movement of visitors in
valleys of displaced villages, specically owners
of cottages and tourists. Seasonal migration could
also be caused by bison preference for open habi-
tats, which are more frequently used during winter
period by both sexes. However, bison rarely move
further than 5 km from the forest edge (Kowalczyk
et al. 2013). The valleys with displaced villages in
Poloniny achieve these conditions.
In general, eastern slopes were the most occu-
pied (34.7%), followed by western (28.0%), southern
(22.0%) and northern slopes (15.3%). Southern and
western slopes at higher altitudes from 620 to 650
m a.s.l. were commonly occupied by bison during
summer. Small incidence on northern slopes may
be caused by the coldest climatic conditions and
consecutive inuence to composition of ora. On
the other hand, incidence on slopes with eastern,
southern and western exposures may be caused
by long-lasting sunshine and stronger solar radia-
tion. The quality of winter refuges depends on the
amount of canopy cover and the dominant tree spe-
cies (Wołoszyn-Gałęza et al. 2016). In winter, bison
remained in displaced villages and fed mainly on
apple orchards similarly to the reintroduced Euro-
pean bison in the Bieszczady Mountains (the East-
ern Carpathians, Poland), which actively selected
broken canopy forests with a high density of un-
derstory vegetation (Wołoszyn-Gałęza et al. 2016).
The European bison is a gregarious animal.
The most observed units are small bull groups and
groups composed of cows, calves and bulls. In this
second type of group cows, calves aged 2-3 years
and sometimes adult bulls were observed (Krasiński
and Krasińska 1992, 1994; Krasiński et al. 1994b,
1999; Pucek et al. 2004). Groups of European bison
are not family units and changes in structure and
size of groups are inuenced by other factors. Sea-
sonal factors include bulls joining new groups in the
rutting season and calves being introduced follow-
ing calving. Others changes are a result of behav-
ioural factors. Individuals move between groups at
frequent group meets. Switching between groups is
most common in young bulls (Krasińska et al. 1987).
Environmental factors inuence the average size of
these groups. In different populations, groups are
most often comprised of an average of 8-13 animals
(Krasiński and Krasińska 1992, 1994; Krasiński et
al. 1994b, 1999; Pucek et al. 2004) or 6-20 animals
(Pčola and Adamec 2007). Occasionally, groups of
2-92 animals were observed in the Białowieża For-
est, but more frequently groups of 20 animals (65-
85%) were observed in this location (Krasińska et
al. 1997, Pucek et al. 2004). Small bull groups occur
in all populations and include two animals on aver-
age (1-11). A solitary life is led by more than 50%
of the bulls (Krasiński and Krasińska 1994, Krasiński
et al. 1994a, Pucek et al. 2004). Large mixed groups
of cows, calves and bulls and groups comprised en-
tirely of bulls are observed in winter around feeding
sites. The number of groups of different sizes is af-
fected by the size of population. Mean air tempera-
ture during the winter period and duration of perma-
nent snow cover inuence home ranges of bulls in
winter. Mobility of individuals is limited by low tem-
peratures and long-lasting snow cover (Krasińska et
al. 2000, Pucek et al. 2004). Food supplementation
Table 1. Overview of territory expansion of European
bison in Poloniny National Park. Processed according
to Adamec (2006), Pčola and Adamec (2007), Pčola
and Gurečka (2008).
Discussion
During the reintroduction period in Poloniny
National Park between December 17th, 2004 and
February 17th, 2017, the total area of territory was
106.79 km2. When compared to studies by Adamec
(2006) and Pčola and Adamec (2007), this area grew
from 72.53 km2 to 106.79 km2 (Table 1). Pucek et
al. (2004) state that a satisfactory area for a popu-
lation of 50-70 animals is approximately 200 km2.
According to observations, approximately 25 Euro-
pean bison live in PNP as part of the Slovak herd
within a core area of 106.79 km2 (not including the
Polish herd and Slovak migrating bulls). Compared
to Pčola and Gurečka (2008), the winter area (Octo-
ber to March) was larger, growing from 22.04 km2
to 81.33 km2 (59.29 km2 growth) and the area in-
habited during the vegetation period (April to Sep-
tember) grew from 69.13 km2 to 104.27 km2 (35.14
km2 growth). Boundaries of total area were reached
mostly in vegetation season, rst in the period after
reintroduction (December 10th, 2004 to December
31st, 2007). During other periods, bison inhabited
the core area. This data conrms the conservation
of European bison territory in PNP.
The individuals from Slovak herd were recorded
several times at the border with Poland that runs
along the ridge of the Poloniny Mountains. On the
other side of the border are the Bieszczady Moun-
tains, where the Polish population of European bison
lives (Wołoszyn-Gałęza et al. 2016). Slovak individu-
als have repeatedly moved into the border area and
might possibly get across the border into the Pol-
ish territory, but this behaviour was not recorded,
because observers did not collected transboundary
data. It can be assumed that potential interconnec-
tions with Polish herd could be near Hostovický creek
and along the intersection of the boundary with the
territory of the bison from Slovak area (Fig. 2).
Total average altitude for the year is 501 m a.s.l.
Bison inhabit the lowest altitudes in winter, fol-
lowed by spring. During autumn and summer ani-
mals inhabit the highest altitudes. Winter incidence
at lower altitudes is inuenced by the availability
of supplementary food in hay feeders, long-last-
ing snow cover, low temperatures and decreased
movements of visitors due to winter conditions. Oc-
currence at higher altitudes during vegetation pe-
riod, specically summer, may be due to a sufcient
Period Total
area
(km2)
Winter
area
(km2)
Vegeta-
tion pe-
riod area
(km2)
10 December 2004
to 2 February 2006
72.53 No data No data
10 December 2004
to 3 August 2008
78.24 22.04 69.13
10 December 2004
to 17 February 2017
106.79 81.33 104.27
4
M. Bielik, M.
Janiga & Z.
Ballová
5
Spatio-temporal
distribution of the
European bison in
the Poloniny Na-
tional Park
can also be responsible for their low mobility and
staying in valleys during winter (Krasińska et al.
2000). We found that during the winter period, bi-
son remain close to the feeders when supplied with
hay. However, supplementary feeding alone cannot
represent a sustainable solution, because it causes
bison aggregation and may lead to higher parasitic
transmission (Ramos et al. 2016). Individuals wander
the forest constantly due to its high claim to food. In
the rst year of life, calves eat 8.5 kg per day, while
young animals (2-3 years) consume19.5-28.5 kg per
day, and adults eat 23-32 kg of fresh food per day.
40% of this food is composed of leaves and sprouts
(Gębczyńska and Krasińska 1972, Pucek et al. 2004).
Feeding habits of European bison were researched
mostly in Cejjskijj zakaznik in Russia, Białowieża
Forest (Poland and Belarus) and Prioksko-Terrasnyjj
reserve in Russia (Pucek et al. 2004). 131 plant spe-
cies, with 27 species of trees and shrubs, 14 species
of grasses and sedges and 96 species of dicotyledon-
ous plants were described in the study of Borowski
and Kossak (1972) as part of the bison diet in the
Polish part of Białowieża Forest. Herbs, grasses and
sedges represent 67% and trees and shrubs 33%
of the whole diet. All wild-living groups formed in
Białowieża Forest in winter are fed with supplemen-
tary food, specically with hay (Krasiński 1978, 1983;
Krasiński and Krasińska 1994; Pucek et al. 2004).
European bison went extinct in the wild until
populations were restored during the 20th century.
Today, the species is still listed by the International
Union for Conservation of Nature (IUCN Red List
of Threatened Species) as a species vulnerable to
extinction. Despite of the increasing number of
European bison, there are some specic problems
regarding their sustainability in human inhabited
areas (Hofman-Kamińska and Kowalczyk 2012),
low genetic diversity (Oleński et al. 2018) and suf-
cient size of their territory with adequate habitat
connectivity (Ziółkowska et al. 2016). Expansion of
European bison populations is necessary, and ana-
lysing long-term data about bison space utilization,
habits and movement patterns in existing popula-
tions may help to achieve this goal.
The European bison, as an extremely inbred
extinct species requires effective management to
sustain maximum genetic diversity (Oleński et al.
2018). Our results provide better understanding of
bison space utilization and location of natural mi-
gratory routes during the period of their reintroduc-
tion to the Poloniny NP. The ndings obtained in
this study are benecial for sustainable manage-
ment of the European bison and for facilitate and
subsequently maintenance migration of bison be-
tween neighbouring trans-border herds.
Acknowledgements
We would like to thank the Ministry of Education,
Science, Research and Sport of the Slovak Republic
for the grant from the European Structural Funds to
our project. The present study has been funded by
the ITMS (Project numbers: 26210120006). For Eng-
lish language editing and proofreading, we are in-
debted to Amanda Clarahan. We are grateful to Mar-
tina Vlasáková and Ján Lempeľ for advices and help
with sample collection, to Mário Perinaj for his com-
plaisance, and to Jaroslav Solár and Vladimír Hurta
for help with data analysis.
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Received 8 August 2018; accepted 6 November 2018.
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Janiga & Z.
Ballová
