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DOI: ./conl.
LETTER
Testing a conservation compromise: No evidence that public
wolf hunting in Slovakia reduced livestock losses
Miroslav Kutal1,2Martin Duľa1,2Alisa Royer Selivanova1
José Vicente López-Bao3
Faculty of Forestry and Wood
Technology, Department of Forest
Ecology, Mendel University in Brno,
Brno, Czech Republic
Carnivore Conservation Programme,
Friends of the Earth Czech Republic,
Olomouc, Czech Republic
Biodiversity Research Institute, CSIC,
Oviedo University, Principality of
Asturias, Oviedo University, Mieres,
Spain
Correspondence
Miroslav Kutal, Faculty of Forestry and
Wood Technology, Department of Forest
Ecology, Mendel University in Brno,
Zemědělská , Brno, Czech
Republic. Email:
miroslav.kutal@hnutiduha.cz
Funding information
Regional Government of Asturias
(GRUPIN research grant), Grant/Award
Number: AYUD//; Spanish
Ministry of Economy, Industry and
Competitiveness, Grant/Award Number:
CGL--R AEI/FEDER EU
Abstract
Variation in the legal status and management of wolves (Canis lupus) across EU
Member States provides a good opportunity to test the effectiveness of different
practices to reduce livestock losses. This opportunity for testing is particularly
useful for lethal interventions, as they are among the most controversial actions
within the large carnivore management toolbox. We aimed to test a conservation
compromise adopted in Slovakia, based on a public wolf-hunting scheme and
annual hunting quotas between and , and partially justified to reduce
livestock losses. We assessed whether this hunting scheme influenced livestock
depredation levels (at the district level). Wolves in the area fed mainly on wild
ungulates (.% of consumed biomass). While domestic sheep comprised only
.% of the diet, they were dominant among the reported livestock killed by
wolves (.%). Using two different approaches, we did not observe a relation-
ship between the number of killed wolves and livestock losses. Alternatively, a
negative relationship between wild prey biomass and livestock losses was found.
Since , public wolf hunting has not been conducted in Slovakia, and there is
no merit in the previous justification for this conservation compromise to reduce
livestock losses.
KEYWORDS
Canis lupus, evidence-informed conservation, large carnivores, livestock depredation, wolf
hunting, wolf management
1 INTRODUCTION
Large carnivore conservation is challenging in human-
dominated landscapes, where the compatibility of their
presence with livestock farming (particularly extensive
practices) represents one of the oldest conflict drivers in
the conservation of these species (Lute et al., ). From
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the
original work is properly cited.
© The Authors. Conservation Letters published by Wiley Periodicals LLC.
the perspective of either wildlife management or live-
stock welfare (López-Bao & Mateo-Tomás, ), a range
of different nonlethal and lethal interventions have been
proposed, and used, to minimize the risk of livestock
depredations (e.g., Eklund et al., ; Lorand et al., ;
van Eeden et al., ). Balancing between large carnivore
conservation and farming or hunting interests is politically
Conservation Letters. ;e. wileyonlinelibrary.com/journal/conl 1of8
https://doi.org/./conl.
2of8 KUTAL .
desirable, which often results in controversial conservation
compromises.
Lethal interventions are among the most controversial
issues when dealing with large carnivores (Lute et al.,
) and currently receive less public support (Lute &
Attari, ) and increased attention due to several ethi-
cal concerns (Vucetich & Nelson, ). Furthermore, four
independent reviews published between and
(van Eeden et al., ) agreed on the need to improve stan-
dards of evidence used in evaluating interventions against
carnivore attacks and highlighted how these standards
have thus far been much lower for lethal compared to non-
lethal interventions (see also Lorand et al., ;Treves
et al., ), although there are also relatively few robust
studies evaluating the latter (see Eklund et al., ).
Variation in the wolf’s legal status and management
measures implemented among EU Member States, states
of the United States or Canadian provinces provides useful
opportunities to test the impact of different wolf man-
agement approaches (e.g., Treves et al., ) on conflict
mitigation and wolf conservation. Wolf public hunting
schemes are frequently justified as a tool to reduce live-
stock depredations and facilitate safe coexistence with
humans (e.g., Vucetich et al., ). Apart from the diverse
legal, ethical, and ecological considerations, lethal inter-
ventions in this case should only be considered an effective
tool if there is an observable reduction in livestock losses
after their implementation that does not come at the cost
of threatening the viability of wolf populations.
Interestingly, the relationship between wolf public hunt-
ing schemes and the dynamics of livestock losses has
received less research attention than expected consider-
ing the remarkable public interest around this topic (e.g.,
Delibes-Mateos, ). Standards for available studies vary
considerably, but excluding studies with flawed design still
shows weak or uncertain effects of lethal control (Treves
et al., ). Previous studies that have explored the rela-
tionship between lethal control (mainly carried out by
agency authorities; i.e., culling) and livestock depredation
have offered contrasting results (e.g., Bradley et al., ;
DeCesare et al., ; Santiago-Avila et al., ), even
when using the same dataset (Kompaniyets & Evans, ;
Poudyal et al., ; Wielgus & Peebles, )andbeing
complex to synthesize a general and unique effect (Grente,
).
In Europe, the Bern Convention of and the Habi-
tats Directive of are the fundamental legal frameworks
for wolf conservation. Wolves are listed in Annex IV of
the Directive (species of community interest in need of
strict protection) in most EU Member States, with some
populations (entire or partial) being listed in Annex V
(species of community interest whose taking in the wild
and exploitation may be subject to management measures,
including regulated hunting). Under either of these
annexes, Member States are required to ensure that wolf
populations reach or maintain a “favorable conservation
status” (Epstein et al., ). Several Member States with a
historical tradition of public wolf hunting have banned this
practice even with the species listed in Annex V. Poland
passed the full protection of wolves into national law in
(Nowak & Mysłajek, ). Public hunting of wolves
has been banned in Romania since (Popescu et al.,
), in Slovenia since (Ferraro & Bombieri, ),
and Spain since (Instruction No. TED// of the
Ministry for the Ecological Transition). In Slovakia, the
wolf protection status has changed substantially after ,
following an infringement procedure at the EU level. The
number of killed wolves dropped by % (Kutal et al.,
and this study), and the species was protected year-round
in a strictly defined area along the borders with the Czech
Republic, Poland and Hungary. This partial protection was
a conservation compromise among the interests of diverse
stakeholders. The Ministry of the Environment of the Slo-
vak Republic listed the wolf as a fully protected species
under the implementation of the Decree no. /.
Although public wolf hunting continues in other Annex
V,orevenAnnex IV, countries (e.g., Sweden; Epstein
et al., ), there is limited knowledge on how public
wolf-hunting schemes influence livestock depredations in
Europe (Krofel et al., ) (see DeCesare et al., for
North America). Such evidence is of paramount impor-
tance as the recent recovery of wolves across EU Member
States, excluding islands (Boitani et al., ; Chapron
et al., ) has intensified the debate on the liberalization
of strict wolf protection due to the social conflicts arising
from livestock depredation (e.g., Kiffner et al., ). Com-
pared to North America, domestic ungulates are a more
common food source for wolves in Europe (Newsome et al.,
;Singeretal.,).
Here, we aimed to test the influence of this conservation
compromise, the limited public wolf-hunting scheme used
in the Slovak Republic between and (i.e., a regu-
lated hunting of nonspecific individuals by private citizens,
based on annual hunting quotas), on livestock depreda-
tion levels at the Slovakian district level. Wolves have long
been hunted in the area without a robust evaluation of
the impact of this practice on wolf population dynamics
or livestock losses (Kutal & Dula, ). Each year, public
wolf hunting was allowed from November until January
, the next year. Quotas were set prior to November each
year on a national or regional level by a special working
group, based on suggestions from district hunting admin-
istrations and from several sources of information such as
official livestock depredation data and results of wolf mon-
itoring (Antal et al., ). However, no methodology has
been developed on the use of livestock depredation data for
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KUTAL . 3of8
FIGURE 1 Study area for analysis of wolf livestock depredation in Slovakia ( districts with reported wolf presence, –, red
line) and model area for detailed analysis of species feeding ecology ( districts, green line). Area-specific data showing the number of sheep
killed by wolves (shades of red) and the number of wolves killed the previous winter in Slovakia (circles).
quota setting, and quotes were set only at the regional level
rather than the district level.
2METHODS
2.1 Data collection
We used information at the level of Slovakian districts on
livestock damages, number of wolves hunted, and wolf
diet. We selected districts (, km; % of total Slo-
vak area) where wolf presence was reported by hunters
during the study period to study the effect of public wolf
hunting on patterns of livestock damage, and districts
( km; .% of area with reported presence of wolves)
in north-western Slovakia to study wolf diet (Figure ).
This region has permanent wolf occurrence and breeding
(Kutal et al., ) and unlike other areas in Slovakia it has
not yet been studied regarding wolf feeding habits (e.g.,
Guimarães et al., ). The district level was the smallest
administrative unit available across all data sources (aver-
age of ± km;− km)andwasonaverage
larger than the size of wolf territories in Slovakia or neigh-
boring Poland (Finďo & Chovancová, ; Jędrzejewski
et al., ).
Overall, sheep is the most common livestock species
in all selected districts (, heads; .% of total
farmed animals), followed by cattle (,; .%) and
goats (,; %; Central Evidence of Livestock, https://
www.cehz.sk/index.jsp). According to the National Forest
Centre database “Polovstat” (https://gis.nlcsk.org/ibulh/
PolovStat/PolovStat), the assemblage of wild ungulates
here consists of wild boar (Sus scrofa, .% of hunted
ungulates), red deer (Cervus elaphus, .%), roe deer
(Capreolus capreolus, .%), fallow deer (Dama dama,
.%), and mouflon (Ovis aries, .%). Slovakian authorities
reported – wolves for the entire country in to
the European Commission, according to Article report-
ing of the Habitats Directive (Černecký et al., ;Eionet
Portal, ). In , around wolves were estimated
in the country, including transboundary animals (Boitani
et al., ).
For the period –, we obtained yearly data on
livestock heads killed by wolves ( heads in total),
estimated the total number of wolves, counted the num-
ber of wolves killed by district in the previous season
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4of8 KUTAL .
( wolves were hunted during the study period; annual
average ±.), estimated the number of wild ungu-
lates, and obtained the number of farmed livestock (State
Nature Conservancy of the Slovak Republic, National
Forest Centre, and Central Evidence of Livestock, respec-
tively). Sheep was the most common livestock type killed
by wolves (.% of livestock heads). Goats and cattle were
reported rather rarely (reaching .% and .% of live-
stock killed by wolves, respectively). These figures account
for .% of sheep, .% of goats, and .% of cattle
of the livestock census. Data on livestock damages older
than were not available at the district level. We
estimated the available biomass of wild ungulates based
on a reverse calculation method, following Kutal et al.
(). We also analyzed wolf scats collected between
and in north-western Slovakia. Details on
scat collection, determination and analysis are described
in Appendix S..
2.2 The relationship between public
hunting and livestock depredations
We adopted two different approaches to explore the effect
of public wolf hunting on livestock depredation levels.
First, we built generalized mixed models using a negative
binomial distribution for the response variable: number of
livestock killed by wolves at the district level. We tested
whether the number of livestock killed in a given year and
district was influenced by the number of wolves shot in
the previous hunting season (i.e., from November of the
previous year until January of that year). We used two
proxies for measuring the impact of public wolf hunting:
(i) absolute number of wolves hunted and (ii) the propor-
tion of wolves hunted from the total estimated number of
wolves at the district level. We considered two additional
competing models in our analyses: (iii) considering the
number of farmed livestock and (iv) considering the esti-
mated biomass of wild ungulates. See Appendix S. for
details of the statistical analysis performed. Second, we ret-
rospectively selected a subset of pairs of consecutive years
(i.e., cases) to run a × before-after control-impact design
(BACI design; Appendix S.). We compared the number
of livestock depredations one year before and after win-
tertime, the period of wolf-hunting season (November to
January ), in a number of treatment and control cases
with and without killed wolves (treatment = cases, con-
trol = cases). See Appendix S. for the criteria used to
select cases for the BACI design, and the statistical analysis
performed.
3RESULTS
3.1 The influence of public wolf
hunting on livestock losses
There were no changes over time in the number of live-
stock losses (rS=−., p=.), number of wolves
killed in the previous season (rS=−., p=.), or
number of districts with reported wolves during the study
period (i.e., wolf range; rS=., p=.). However,
the estimated biomass density of wild ungulates increased
significantly over time (rS=., p=.). Livestock
losses in a given year did not correlate with the num-
ber of wolves hunted in the immediate previous hunting
season (Appendix S), either in absolute (posterior param-
eter estimate: . ±., % BCI: −./.) or relative
terms (posterior parameter estimate: −. ±., %
BCI: −./.). Our × BACI design approach com-
plemented this finding as we did not detect differences in
the number of livestock losses at the district level between
treatment and control groups (Appendix S.).
3.2 Other factors affecting livestock
losses
Wild ungulates dominated the wolf diet (.% of con-
sumed biomass), and cervids were consumed far more
than wild boars (% deer vs. .% wild boar). According
to hunting bags in north-western Slovakia, the commu-
nities of wild ungulates were dominated by wild boar,
followed by red deer and roe deer. We observed a high
selection for roe deer (D=.), a high avoidance of wild
boar (D=−.), and predation on red deer according
to its availability (D=−.; Figure ). Sheep was esti-
mated to account for . % of consumed biomass, and
no other domestic animal was detected in this dataset
(Table ). In this context, the model including the esti-
mated biomass density of wild ungulates was the most
parsimonious model explaining livestock losses in this
area, with a negative influence on the magnitude of losses
(Appendix S).
4DISCUSSION
Based on the two different approaches used in this
study, public wolf-hunting schemes did not correlate with
livestock depredation levels at the Slovak district level.
Although livestock damages in the previous two years
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KUTAL . 5of8
TABLE 1 Diet composition of wolves in the study area of north-western Slovakia (–). Subtotals and totals are marked in bold.
Prey item
Frequency of occurrence
(%)
Biomass consumed
(%) Biomass consumed (g)
Undetermined cervids . . ,
Roe deer Capreolus capreolus . . ,
Red deer Cervus elaphus . . ,
Wild boar Sus scrofa . . ,
Wild ungulates total 97.69 98.86 359,792
Brown hare Lepus europaeus . .
Medium mammals total 0.38 0.52 1898
Bank vole Myodes glareolus . ≤.
Undetermined weasel Mustela spp. . ≤.
Undetermined Soricidae . ≤.
Undetermined small mammal . ≤.
Small mammals total 1.53 0.07 237
Sheep . .
Livestock total . 0.55 2015
Plant material 7.3 ──
Number of scats 260
were used to set the baseline for establishing hunting
quotas, according to the Slovak wolf management plan
(Antal et al., ), the quota system was set on national
and regional scales yearly and could not target specific
hotspots with increased damages. The absence of uniform
procedures for using livestock depredation data through-
out Slovakia to prioritize conflict areas was likely one of
the significant shortcomings of quota setting. The observed
failure in reducing livestock losses is expected if predators
are nonselectively removed (Lennox et al., ). Bradley
et al. () found that even partial removal of a wolf pack
was not effective in reducing livestock depredations if con-
ducted more than days after the depredation event, and
only a marginally significant difference between partial
pack removal and no action was observed if conducted
after days from the depredation (but see Santiago-Avila
et al., ). The assessed public hunting scheme did not
fulfill the criteria for this short-term targeted removal. Two
other studies performed in Montana, USA and Slovenia
found no evidence that removing wolves through public
harvest affected the year-to-year presence or absence of
livestock losses by wolves (DeCesare et al., ;Krofel
et al., ). A small effect (. fewer depredation events
year) was found only within the subset of districts with
conflicts, where public harvest of a greater proportion of
the known wolves in a district reduced the number of
depredations (DeCesare et al., ). Additionally, previ-
ous research has demonstrated that removing wolves from
a particular site did not reduce future risk of recurrence
in wolf depredations in neighboring areas (Santiago-Avila
et al., ). The before-after control-impact design used
in this study does not provide the strongest inference
about the effects of predator control and the use of higher
standards is recommended to address this question more
thoroughly (Treves et al., ). For example, some finer
effects of wolf culling could also be hidden by the relatively
coarse scale of our study at the district level (Santiago-Avila
et al., ).
The overall results of diet analysis closely resemble
observations of the dietary preference of wolves in Central
Europe, where wild ungulates were by far the dominant
prey item and livestock was marginally represented (e.g.,
Guimarães et al., ; Nowak et al., ). The lack of exact
data on the abundance of prey communities, which were
not studied by robust methods in most of referred studies
on wolf diet, could cause a possible bias in prey selection.
The lack of a relationship between wolves hunted and
livestock killed by wolves could also be caused by negli-
gible representation of livestock in the diet of Slovakian
wolves found in north-western Slovakia within the same
period (this study) or in central and eastern Slovakia in
– (Guimarães et al., ). In addition to the cor-
rect implementation of effective livestock damage preven-
tion measures (to reduce the vulnerability of livestock to
wolf attacks), the availability of wild ungulates is an impor-
tant factor shaping wolf diet patterns in human-dominated
landscapes (e.g., Mayer et al., ). Our analysis indicated
that abundant wild prey may actually avert wolf predation
on livestock, but these conclusions should be interpreted
with caution—a large area with permanent wolf presence
has lower biomass density due to higher altitudes and
overall lower productivity of mountain habitats and the
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6of8 KUTAL .
FIGURE 2 Composition of wild ungulate communities based
on hunting bag, wolf prey, and preference/avoidance of wolves
based on Ivlev’s electivity index in study area of north-western
Slovakia (–).
densities of ungulate populations cannot be augmented
due to generally high damages from deer browsing in the
forest (Finďo et al., ). Data on other important factors
which drive livestock vulnerability, such as presence and
effectiveness of damage prevention measures, are miss-
ing for each reported case in official statistics. We also
acknowledge that reported damages on livestock may not
be complete, since some minor losses may not be discov-
ered or reported due to bureaucratic reasons. However,
considering the average annual number of damages was
livestock heads and depredations were one of basis for
setting hunting quotas, we expect that the proportion of
missing head was low. We used the only available official
data and believe they are not significantly underestimated
due to the opportunity for financial compensation.
Low levels of livestock predation by wolves in our study
and in other studies from Central Europe indicate that
wolves in this area do not rely on livestock for their per-
sistence. Therefore, the argument for public hunting of
wolves to prevent severe sheep losses and protect food
security in Slovakia (Kutal & Dula, ) lacks evidence.
Wolves received year-round protection in Slovakia in ,
making it the last country in Central Europe to include
year-round protection of the species under national law.
There are strong political incentives to scapegoat large car-
nivores (Chapron & López-Bao, ), and new attempts
for lethal management of wolves in Europe may arise
in the near future using similar arguments. Whether
the conflict in Slovakia around wolves was mitigated
though this public hunting scheme, despite no observ-
able effects on livestock depredations, cannot be answered
if no human-dimensions research took place before and
after the implementation of this conservation compro-
mise. However, available studies from other regions did
not provide much evidence that attitudes towards wolves
become more positive after legal wolf hunting (Browne-
Nuñez et al., ), or that public wolf-hunting schemes
will decrease illegal killing of the species (e.g., Chapron &
Treves, ; Suutarinen & Kojola, ).
Nonlethal methods such as livestock guarding dogs or
fences have been found effective in livestock protection
(see review in van Eeden et al., ), although only few
studies have had high quality experimental design (Eklund
et al., ). Yet, there was no relevant mechanism for
compensating the increased expenditure on preventive
measures by public funds in Slovakia, which can economi-
cally burden livestock breeders in higher-risk areas (Kutal
&Dula,), even when preventing wildlife attacks on
livestock is also a matter of animal welfare, included in
EU regulations on livestock welfare (López-Bao & Mateo-
Tom ás, ). Therefore, we urge for the generalized use
and support for the effective implementation and mainte-
nance of nonlethal interventions (Frank & Eklund, ),
and for efforts in increasing acceptance of the use of these
measures among end-users (Eklund et al., ) in order to
facilitate the coexistence of wolves and livestock farmers in
this cultural landscape.
AUTHOR CONTRIBUTIONS
MK and JVLB designed the study. MK led the writing of
the manuscript. MK, MD, and ARS collected and analyzed
data. JVLB and MK ran the analyses. All authors reviewed
and commented on the paper.
ACKNOWLEDGMENTS
We are grateful to the dedicated volunteers of Wolf Patrols
and staff of PLA Kysuce for their participation in field work
and to the State Nature Conservancy of the Slovak Republic
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KUTAL . 7of8
and National Forest Centre for providing data on livestock
damages and hunting counts. We also acknowledge com-
ments of four anonymous reviewers that helped to increase
the quality of the manuscript.
CONFLICT OF INTEREST STATEMENT
The authors declare no conflicts of interest.
DATA AVAILABILITY STATEMENT
The data that support the findings of this study are
openly available at https://zenodo.org/doi/./zenodo.
.
ETHICS STATEMENT
The search for and sampling of scats involved noninvasive
methods did not affect the studied animals.
ORCID
Miroslav Kutal https://orcid.org/---
Martin Duľa https://orcid.org/---
Alisa Royer Selivanova https://orcid.org/--
-X
José Vicente López-Bao https://orcid.org/--
-X
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SUPPORTING INFORMATION
Additional supporting information can be found online
in the Supporting Information section at the end of this
article.
How to cite this article: Kutal, M., Duľa, M.,
Selivanova, A. R., & López-Bao, J. V. (). Testing
a conservation compromise: No evidence that
public wolf hunting in Slovakia reduced livestock
losses. Conservation Letters, e.
https://doi.org/./conl.
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