Journal for Nature Conservation 47 (2019) 28–37

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Journal for Nature Conservation

journal homepage: www.elsevier.com/locate/jnc

The change in the attitudes of Czech hunters towards Eurasian lynx: Is T

poaching restricting lynx population growth?
⁎,1
J. Červenýa,1, J. Krojerová-Prokešováb,c, , T. Kuštaa, P. Koubeka,b
a
Department of Forest Protection and Game Management, Faculty of Forestry and Wood Sciences, Czech University of Life Sciences Prague, Kamýcká 1176, 165 21 Prague
6, Suchdol, Czech Republic
b
Institute of Vertebrate Biology of the Czech Academy of Sciences, Květná 8, 603 65 Brno, Czech Republic
c
Department of Zoology, Fisheries, Hydrobiology and Apiculture, Faculty of AgriSciences, Mendel University in Brno, Zemědělská 1, 613 00 Brno, Czech Republic

A R T I C LE I N FO A B S T R A C T

Keywords: Even though the Eurasian lynx is a less controversial species than the wolf or brown bear, poaching remains a
Anonymous survey major cause of lynx mortality in Europe, potentially threatening population expansion in key areas. Our study
Conservation was designed to explore the attitude of hunters and other stakeholders (students of secondary forestry schools
Hunting and University forestry faculties) towards lynx and their experience with illegal killing of lynx in the Czech
Perception
Republic. Self-administered questionnaires were addressed to local hunters and to students in 2001 and again in
Poaching
Questionnaires
2015. The survey in 2001 was conducted in two separate hunting regions of Bohemia, one where lynx have been
artificially reintroduced and another in which lynx have become established through natural colonisation. In
2015 the survey was extended to two further areas where lynx have re-established themselves through natural
colonisation, in the east of the Czech Republic. Altogether 415 and 922 questionnaires were completed in 2001
and 2015, respectively. The attitude of hunters towards lynx was not affected by hunting region, lynx population
density or nature of the population (human re-introductions vs natural recolonisation), but attitudes became
more negative in 2015 than they had been in 2001. The majority of hunters still believed that lynx had negative
effects on other wildlife and 27% stated that they do not wish to co-exist with lynx. Half of secondary school
students and a third of tertiary students still believe that lynx threaten roe deer stocks, even though roe and red
deer are present at high density and cause extensive damage in commercial forestry. Hunters as well as forestry
students had first-hand knowledge about illegal hunting of lynx. The proportion of hunters admitting to having
poached lynx themselves was 10% with the proportion of repeated illegal kills made by the same person in-
creasing between 2001 and 2015. Population modelling suggested that at least 25% of the population might be
poached annually, sufficient to restrict population growth and further expansion of lynx distribution.

1. Introduction competition for game (Røskaft, Händel, Bjerke, & Kaltenborn, 2007;
Baker, Boitani, Harris, Saunders, & White, 2008; Røskaft, Bjerke,
Despite the fact that Europe is densely populated and landscape is Kaltenborn, Linnell, & Andersen, 2003). Negative attitudes can in turn
heavily fragmented, a number of species of large carnivores have shown influence human behaviour, including poaching (illegal killing) which
extensive recovery in recent decades (Chapron et al., 2014). This is considered by many authors to be the main threat for large carnivore
comeback is due to a combination of favourable legislation and suc- populations in Europe (Andrén et al., 2006; Breitenmoser et al., 2000,
cessful reintroductions, changes in human land-use and adaptation of 2010; Hell, Slamečka, & Gašparík, 2004; Kowalczyk, Gorny, & Schmidt,
large carnivore populations to human pressure (López-Bao, Kaczensky, 2015; Liberg et al., 2012; Linnell, Breitenmoser, Breitenmoser-Würsten,
Linnell, Boitani, & Chapron, 2015; van Heel, Boerboom, Fliervoet, Odden, & von Arx, 2009; Mykrä, Pohja-Mykrä, & Vuorisalo, 2017;
Lenders, & van den Born, 2017). Although public support for large Okarma et al., 2002).
carnivore protection seems to be growing, some stakeholder groups still There has been a remarkable increase in the number of studies ex-
have a negative attitude towards large carnivores, arising mainly out of amining people’s attitudes towards large carnivores in Europe and
depredation of domestic animals, fear for personal safety and perceived North America over the last two decades. To date, most research has

⁎
Corresponding author at: Institute of Vertebrate Biology of the Czech Academy of Sciences, Květná 8, 603 65 Brno, Czech Republic.
E-mail address: krojerova@ivb.cz (J. Krojerová-Prokešová).
1
These authors contributed equally to this work.

https://doi.org/10.1016/j.jnc.2018.11.002
Received 27 July 2018; Received in revised form 19 October 2018; Accepted 9 November 2018
1617-1381/ © 2018 Elsevier GmbH. All rights reserved.
J. Červený et al. Journal for Nature Conservation 47 (2019) 28–37

been conducted on the more controversial species such as wolves and Koubek & Červený, 1996). However, after the initial rapid increase,
brown bears (e.g. Bjerke, Skogen, & Kaltenborn, 2002; Williams, numbers declined and population has stagnated during the last two
Ericsson, & Heberlein, 2002; Ericsson & Heberlein, 2003; Majić, Taussig decades (Wölfl et al., 2015; Heurich et al., 2018). At the turn of 20th
de Bodonia, Huber, & Bunnefeld, 2011; Treves, Naughton-Treves, & and 21th century lynx also reoccupied the northern part of Bohemia by
Shelley, 2013; Browne-Nuńez, Treves, MacFarland, Voyles, & Turng, natural recolonization from the population established in the south-
2015) with fewer studies focusing on Eurasian lynx Lynx lynx (Bath, west; at present, lynx occurs there only occasionally.
Olszanska, & Okarma, 2008; Červený, Koubek, & Bufka, 2002; The recovery of lynx populations, especially in SW Bohemia, where
Lescureux et al., 2011; Liukkonen, Mykrä, Bisi, & Kurki, 2009). Per- the lynx was artificially reintroduced, was accompanied by a great
ceptions of large carnivores are influenced by people’s relationship with controversy, in part because of predation of domestic livestock (see
nature and by expected or actual personal and/or economic risk at- Kovařík et al. (2014) for details about sheep breeders’ attitude to large
tached to large carnivores (2007, Kleiven, Bjerke, & Kaltenborn, 2004; carnivores in the Czech Republic). However, the damage caused by lynx
Kovařík, Kutal, & Machar, 2014; Naughton-Treves, Grossberg, & Treves, to livestock is relatively small (in comparison to some other species) at
2003; Røskaft et al., 2003; Vittersø, Bjerke, & Kaltenborn, 1999). 1–5 cases per year (Fig. S1) and conflicts with livestock farmers were
Ericsson and Heberlein (2003) highlighted the importance of studying subsequently reduced by implementation of preventive measures and
the attitude of key stakeholders, hunters and livestock farmers, who are financial compensation (Act No. 115/2000 Coll., on compensation of
most directly affected by large carnivores but usually not included in damage caused by selected specially protected animals). At present, a
general public surveys since they form only a minority of the public. greater problem of human-lynx coexistence is the potential for com-
Hunters’ perceptions of large carnivores are described in several petition perceived between lynx and hunters, especially roe deer hun-
studies (Andersone & Ozoliņš, 2004; Bjerke, Reitan, & Kellert, 1998; ters, who are supposed to be the main perpetrators of illegal killing -
Ericsson & Heberlein, 2003; Linnell, Swenson, & Andersen, 2000; they compete with large predators for the same prey and value them as
Lüchtrath & Schraml, 2015; Naughton-Treves et al., 2003; Røskaft trophies, are legal owners of firearms and may be legally armed in the
et al., 2007). Most of these studies report that hunters have negative forests. There is good evidence for illegal killing of lynx within the
attitudes towards large carnivores due to a perceived competition for Czech Republic. During 1995–2003 in southwestern (SW) Bohemia 56
game, especially ungulates (Ericsson & Heberlein, 2003; Linnell et al., skulls from poached lynxes were collected for craniological study and of
2000; Naughton-Treves et al., 2003; Røskaft et al., 2007). It was shown 16 radio-collared lynx being monitored in this area, it is known for
that people in areas with an established tradition of big game hunting certain that three were shot and five more were also mostly probably
are more likely to accept illegal killing of carnivores because they are killed by poachers (Červený, Bufka, & Koubek, 2006; Červený &
competing with them for the same prey (Røskaft et al., 2007; Gangaas, Koubek, 2000; Červený et al., 2002). Further, we have also known of at
Kaltenborn, & Andreassen, 2013). The negative attitude of hunters can least two other cases of lynx shooting within last 5 years in eastern
be also a result of defending their social identity and distinguishing Moravia, where the population has also stagnated (Krojerová,
themselves from other social groups, such as nature conservationists Barančeková, Homolka, & Koubek, 2014). For future success in con-
(Lüchtrath & Schraml, 2015; van Heel et al., 2017). Moreover, there is servation of lynx within a human-dominated landscape, therefore, it is
no doubt that the concept of the hunting of large carnivores themselves crucially important to understand the attitudes of hunters towards
as a quarry species has had a long tradition in Europe and large car- sharing their landscape with an animal viewed as a potential compe-
nivores are still considered to be a valuable game species (Linnell, titor (see also Ericsson & Heberlein, 2003; van Heel et al., 2017).
Swenson, & Andersen, 2001; Liukkonen et al., 2009). In the light of the importance of the hunters’ role for the success of
Available data demonstrate that lynx have a different position in the large carnivore conservation, the aims of our case study were 1) to
public eye, from that taken in relation to other large carnivores such as obtain the knowledge of the hunters’ perception of lynx and to see if the
wolf and bear, being far less known and in general generating less attitude has changed (positively or negatively) during the last 15 years;
polarized views (Lescureux et al., 2011). Lynx as a species is strictly 2) to assess the attitude towards lynx of students of secondary forestry
protected by international and national laws in all Central European schools and of University forestry faculties as a presumably better-
countries. Lynx is listed in Annex II (habitat protection) and Annex IV educated group of respondents, and 3) to evaluate whether the attitude
(strictly protected species) of European Directive 92/43/EEC on the of hunters or students differ between regions where lynx have re-es-
conservation of natural habitats and of wild fauna and flora (the Ha- tablished themselves by natural colonisation and those where lynx
bitats and Species Directive). The Directive provides a robust frame- population had been established by artificial reintroduction.
work for achieving favourable conservation status of the species in
Europe. Despite this, illegal killing by hunters remains a major cause of
lynx mortality in Europe. Based on recent general estimates, up to 30% 2. Material and methods
of the European lynx population may be poached annually (Andrén
et al., 2006; Breitenmoser-Würsten et al., 2007; Kowalczyk et al., 2.1. Respondents and anonymous survey
2015).
The lynx was extirpated from the Czech Republic at the turn of 19th In any survey of somewhat sensitive topics, the core concerns of
and 20th century, firstly from the western part of the country - Bohemia respondents relate to potential consequences to themselves of in-
(Šumava National Park) between 1835–1894, then also from the volvement (Hogberg, Treves, Shaw, & Naughton-Treves, 2016; Lee &
eastern part - Moravia (Moravskoslezské Beskydy Mts) between Renzetti, 1990). It has been demonstrated that respondents are more
1912–1928 (Červený & Bufka, 1996; Heurich & Wölfl, 2002; Kratochvíl likely to report sensitive information in self-administered ques-
& Vala, 1968). While the Moravian part of the country was re-occupied tionnaires than in interviews (Tourangeau & Yan, 2007) and when
in 1980s through the natural expansion of the Carpathian lynx popu- anonymity is provided (Murdoch et al., 2014; Ong & Weiss, 2000).
lation (Anděra & Červený, 2009), the recovery of lynx in southwestern Considering our objective to enquire about possible illegal hunting of
Bohemia was as a consequence of human-mediated reintroductions, lynx in the past, which would potentially put respondents at the risk of
firstly in the Bavarian Forest National Park between 1970–1974 self-incrimination, we selected a self-administered anonymous ques-
(Festetics, 1980; Wotschikowsky, 1978), and later on the Czech side in tionnaire survey. A semi-structured questionnaire contained both
the Šumava National Park in 1982–1989 (Nováková, 1997; Trpák, closed and open-ended questions (Fig. S2), and with no questions on
1985). These reintroductions were successful, and the population individual characteristics such as age, sex and education level included.
(currently known as the Bohemian-Bavarian-Austrian (BBA) popula- Specifically, respondents (hunters and forestry students) were
tion) increased in numbers during 1990s (Červený & Bufka, 1996; asked:

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J. Červený et al. Journal for Nature Conservation 47 (2019) 28–37

i) how they view the re-establishment of lynx within the Czech fauna (Fig. 1).
ii) whether or not they consider re-establishment of lynx to have a As well as being circulated among hunters in these different areas,
positive or negative effect on other wildlife in general questionnaires were also addressed in both surveys to students of game
iii) whether or not they considered that hunters’ main quarry (roe deer) management at two forestry faculties and to students of secondary
was negatively affected by lynx predation forestry schools – three situated in SW Bohemia, one in N Bohemia and
iv) if they felt that lynx populations should be actively managed by one in Moravia in both surveys (see Fig. 1, Table 1 for details).
hunting and The significance of changes in the proportion of answers obtained
v) which are the main reasons for illegal killing of lynx for particular questions in 2001 and in 2015 was evaluated using chi-
squared test.
Because only relatively few students would be expected to have Questionnaires for hunters were distributed through personal con-
hunting licences, in the survey of hunters only, the additional question tacts within the hunting community, or with the help of the local au-
was asked: thority for game management or the Czech-Moravian Hunting
Association, the largest hunting association in the Czech Republic (of
• if they have personal experience with illegal killing of lynx which about 90% of hunters are currently registered members). This
helped us to increase cooperation and trust, and subsequently increased
Due to differences in the history of lynx extermination and its re- the response rate within this community. Because it was not possible
covery (whether by natural recolonisation or artificial introduction), accurately to assess how many questionnaires were distributed in either
the attitude of local people in different regions within the Czech survey, the response rate among hunters was calculated as the number
Republic may differ. In 2001 we decided to address an anonymous questionnaires returned as a proportion of all registered hunters and
survey to local hunters in two different regions: (1) southwestern (SW) will thus generally offer an underestimate of actual response rate
Bohemia where lynx populations were re-established by artificial re- (Table 1). Response rates among students were more conventionally
introduction (and with now the highest population density of lynx in calculated as a proportion of returned questionnaires to all distributed
the Czech Republic); (2) northern (N) Bohemia, where lynx had already (Table 1).
become established by natural colonisation by 2001 but as a result of
the previous reintroduction; it is of note that since 2001 the status of 2.2. Population growth modelling
lynx population in N Bohemia has changed from permanent to only
sporadic occurrence. A preliminary report on some findings of this 2001 In order to explore the possibility that illegal killing was inhibiting
survey was presented by Červený et al. (2002). In 2015 we decided to expansion of population numbers and distribution of lynx within the
repeat the survey in both Bohemian areas and to include two other areas studied (which together represent the majority of the distribu-
hunting regions in Moravia (3) the area of eastern (E) Moravia that tional area of lynx within the Czech Republic as a whole), a simple
represents an area of natural recovery of lynx but retains at present a modelling approach was used to simulate the increase in recorded po-
rather lower population density and (4) northern (N) Moravia, where, pulation size from 80 lynx individuals estimated in 2000 (Poledníková
rather as now in north Bohemia, lynxes are reported sporadically et al., 2015; Wölfl et al., 2015) to current population size using the

Fig. 1. The Czech Republic with marked hunting regions, forestry universities and secondary forestry schools were the survey was conducted. The current Eurasian
lynx distribution range (white –permanent occurrence, light grey - sporadic occurrence) according to Chapron et al. (2014) is outlined in EEA grid.

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J. Červený et al. Journal for Nature Conservation 47 (2019) 28–37

Table 1
The numbers of hunters in particular hunting regions and the number of students of secondary forestry schools and forestry universities examined in 2001 and 2015.
stakeholders hunting region/school 2001 2015

possible returned % possible returned %

hunters South-western (SW) Bohemia 11 236 171 2 10 128 184 2

Northern (N) Bohemia 936 33 4 3 852 42 1
Northern (N) Moravia not included 2 835 85 3
Eastern (E) Moravia not included 4 212 94 2
together 12 172 204 2 21 027 405 2
students of secondary forestry schools Secondary Forestry School (SFS) Písek 77 42 55 154 147 95
Secondary Forestry School (SFS) Vimperk 35 30 86 cancelled
Secondary Forestry School (SFS) Žlutice not included 48 22 46
Czech Forestry Academy (CFA) Trutnov 56 30 54 62 59 95
Secondary Forestry School (SFS) Hranice 71 31 44 76 70 92
together 239 133 56 340 298 88
students of forestry faculties Faculty of Forestry and Wood Sciences Czech University of Life Sciences in Prague 115 55 48 151 136 90
(CZU)
Faculty of Forestry and Wood Technologies Mendel University Brno (MENDELU) 60 23 38 129 83 64
together 175 78 45 280 219 78
together 12 586 415 3 21 647 922 4

formula Nt+1 = Nt + n – m, where n is the natality and m is the except for N Bohemia in 2015. Only 22% of hunters assessed the role of
mortality of the population. Based on monitoring data sex ratio was lynx in ecosystem positively or more positively than negatively. Lynx
taken as 1:1 (Breitenmoser-Würsten et al., 2007; Kramer-Schadt, impact on wildlife was considered to be most positive by university
Revilla, & Wiegand, 2005; own data), thus the number of females Nf students; however, again more than 10% of forestry students at uni-
was calculated as Nf = Nt/2. Reproductive rate was calculated pre- versities still perceived the impact of lynx on wildlife as negative.
suming that 2/3 of females reproduce annually and that on average 2 In addition to potential effects on biodiversity in general, it was the
kittens are born per reproducing female (Gaillard, Nilsen, Odden, belief of approximately one half of hunters that the presence of lynx has
Andrén, & Linnell, 2014; Kramer-Schadt et al., 2005; Kutal, 2014; Wölfl a negative impact specifically on populations of game species (roe deer
et al., 2015). Recruitment was then calculated as n = 2/3Nf *2. Mor- stocks planned for particular hunting grounds; Fig. 2C). In 2001, 59% of
tality of the population m is formed by the mortality of juveniles hunters in SW + N Bohemia were of the view that the presence of lynx
(known to be up to 50% within first two years after birth (Jedrzejewski endangers roe deer stocks planned within given hunting grounds; in
et al., 1996; Kramer-Schadt et al., 2005) and by the mortality of adults. 2015 the proportion had fallen to 42% (χ2=12.8039, p < 0.001,
We used four different values of 25%, 30%, 35% and 40% mortality of n=430). In both surveys half of students at secondary schools and
adults in our simulations. Observed population trend was compared to approximately one third of university students shared the same opinion
reported estimates of population size; estimates of likely levels of illegal (Fig. 2C).
killing were based on the presumption that natural mortality and traffic
mortality together have previously been estimated to form up to 15% 3.2. Attitude towards legal hunting of Eurasian lynx
mortality of adults (Andrén et al., 2006; Heurich et al., 2018; Wölfl
et al., 2001). In both surveys the vast majority of hunters (more than 85%)
wanted lynx to be legal quarry, even though only 7% were in favour of
3. Results year-round hunting and thus the majority considered that hunting
should be regulated in some way (temporal or spatial; Fig. 3). Across all
3.1. General attitude of hunters and forestry students towards Eurasian lynx areas, only 4% of hunters agree with (current) year-round protection of
lynx. In SW Bohemia, the area with the most abundant lynx population,
In 2015 only 36% of hunters were in support of accepting Eurasian only 1% of hunters approved the year-round protection of lynx in 2015.
lynx within the Czech fauna without any restrictions (Fig. 2A). Almost Perhaps somewhat surprisingly, students of secondary forestry schools
the same proportion of hunters considered lynx presence should be as well as of forestry faculties shared similar opinions (Fig. 3).
restricted to specific sites, mostly to large protected areas, large forest
stands or to montane areas (Table S1). Overall, some 27% of hunters 3.3. Illegal killing of lynx
were strongly against tolerating the presence of lynx in the Czech Re-
public at all (Fig. 2A). In Bohemia we could compare results obtained in In both surveys large numbers of hunters in all hunting regions were
2015 with those from the previous survey in 2001. In SW Bohemia the aware of specific instances of illegal killing of lynx (the proportion
proportion of hunters who expressed strong opposition to the lynx oc- varying between areas from 26 to 46 %; Fig. 4A). Approximately 20%
currence increased significantly from 9.4% in 2001 to 30.4% in 2015 of students at both types of schools admitted knowledge of lynx
(χ2 = 30.4861, p < 0.001, n=355). In N Bohemia the proportion of poaching.
hunters sharing this opinion did not change - 15% and 12% in 2001 and Moreover, in total, some 10% of hunters admitted that they them-
2015, respectively (χ2 = 0.2731, p = 0.872, n=75). Forestry students selves had killed lynx (Fig. 4B). This proportion was the same within
had a more positive attitude toward lynx than hunters with approxi- both surveys even though the number of surveyed questionnaires
mately half of respondents accepting lynx occurrence with any re- doubled. Illegal hunting was admitted mainly in SW Bohemia (15%),
strictions (Fig. 2A). Nonetheless, approximately 10% of students at both but also in other regions including those with only sporadic lynx oc-
secondary and tertiary colleges considered that the lynx should not be currence. In SW Bohemia and N Bohemia some hunters also admitted
present in the Czech wildlife. that they killed more than 1 lynx (up to 6 individuals). The proportion
Hunters as well as students of secondary forestry schools consider of repeated illegal kills made by the same person increased between
lynx to have a negative impact on other wildlife (Fig. 2B). This attitude 2001 and 2015 in both Bohemian hunting areas. The increase was not
was consistent among all hunting regions and within both surveys significant (χ2=4.7926, p = 0.911, n=430).

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J. Červený et al. Journal for Nature Conservation 47 (2019) 28–37

Fig. 2. The general attitude of hunters and students towards Eurasian lynx in the Czech Republic – answers to following questions: (A) Q1=Do you think that the lynx
should be an integral part of the Czech fauna?; (B) Q2=Which effect has the re-establishment of lynx population on the wildlife in the Czech Republic?; (C) Q3=Does lynx
represent the threat for planned roe deer stocks in your hunting ground or hunting ground you know?.

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J. Červený et al. Journal for Nature Conservation 47 (2019) 28–37

hunting, high lynx density, no compensations for game damage, and/or

simple opposition to state authorities/activities of conservationists
(Fig. 5). Fear of human injury was suggested as a possible reason only in
2001 by two students at secondary schools (Table S2).
Scientific population size estimations based on camera-trapping and
non-invasive genetic sampling indicate population size fluctuation
within the range of 50–80 individuals over the period 2000–2015
(Fig. 6; Bartošová, 2004; Hlaváč, Bartošová, Jaskula, & Strnad, 2014;
Krojerová et al., 2014; Poledníková et al., 2015). Analyses of population
growth using the model described showed that adult mortality at the
level of 35% would act to maintain the population size at stable levels
or result in slight decrease in while mortalities below 30% would
permit population growth (Fig. 6). If current populations are assumed
to be stable or slightly decreasing and we estimate that up to 15% of
this mortality is caused by natural mortality (starvation or diseases) and
by vehicle collisions (see below), such calculations would suggest that
Fig. 3. The attitude of hunters and forestry students towards possibility of legal poaching must result in mortality of at least 20% of the adult popula-
hunting of Eurasian lynx in the Czech Republic (Q4: Should the legal hunting of tion overall (although rates may vary between regions).
lynx be allowed in the Czech Republic?).

The main reason given for such illegal kills was presumed damage to 4. Discussion
game populations, especially roe deer. Trophy hunting was given as the
second most popular reason with a clear increase in 2015. When we 4.1. General attitude of hunters and forestry students towards Eurasian lynx
took into consideration only hunters that admitted lynx poaching,
trophy hunting was the most important reason in 2015 contrary to Despite conservation policies and relevant legislation being in place,
2001, when game damage was stated as the main reason for lynx implementation is substantially influenced by the attitudes of local re-
shooting (χ2=7.7315, p < 0.01, n=42; Fig. 5). Livestock damage, sidents towards the species (Decker & Purdy, 1988; West, Igoe, &
aversion against lynx and failure of punishment (only in 2001) were the Brockington, 2006). Because the presence of lynx is often associated
only other reasons offered by poachers. The increase in the importance with a wide range of social or economic conflicts (Odden et al., 2002;
of trophy hunting is evident also from the survey among non-poachers Zimmermann et al., 2010), there is a need, when developing a con-
(χ2=22.8515, p < 0.001, n=562) and university students servation strategy, to focus on the people who share the landscape with
(χ2=13.9674, p < 0.001, n=278), even though these groups stated this species (Bath, 1996). Livestock farmers and hunters are the most
many other reasons for lynx poaching, e.g. the impossibility of legal relevant for lynx conservation and most likely to have interactive re-
lationships with the lynx. These stakeholders usually have different

Fig. 4. Illegal hunting of Eurasian lynx in the Czech Republic – responses to following questions: (A) Q5=Do you have knowledge about the specific case of lynx poaching
in the Czech Republic? (B) Q6=Did you shoot the lynx in the Czech Republic?.

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Fig. 5. Main reasons for illegal killing of lynx given by hunters and students in the surveys in 2001 and in 2015 (Q7 in Fig. S2).

support for wildlife conservation, including large carnivores (Krajhanzl

et al., 2015). In this wider survey, further expansion of lynx distribution
was supported by 76% of 1 857 respondents, by 37% of them strongly.
Only 3% of respondents supported the reduction of current lynx dis-
tribution. Contrary to this survey, our results showed a rather negative
perception by both hunters and (informed) students throughout, plus a
shift towards an even more negative view of lynx. In SW Bohemia, the
area with the largest lynx population, we recorded a significant increase
from 2001 to 2015 in the proportion of hunters who do not want to
coexist with lynx (from 9% to 30%; Fig. 2C), despite large campaign to
raise awareness among different stakeholders including hunters (e.g.
Kutal & Bláha, 2008). This is in accordance with other studies con-
firming that the proportion of people with positive attitudes toward
large carnivores is consecutively decreasing with the duration of their
coexistence (Bisi, Kurki, Svensberg, & Liukkonen, 2007; Dressel,
Fig. 6. Scientific (Poledníková et al., 2015; Wölfl et al., 2015) and hunting Sandström, & Ericsson, 2015; Ericsson & Heberlein, 2003; Røskaft et al.,
(Hunting statistics 2000-2015Hunting statistics 2000-2015) census estimates of 2007) and/or with the increase in their abundance (Bjerke et al., 1998;
lynx population size from 2000 till 2015 and population growth modelling Eriksson, Sandström, & Ericsson, 2015; Røskaft et al., 2007).
curves with different presumed mortality rates of adults (see Material and It has been suggested that hunters had a more positive response
methods). towards lynx in areas where populations had shown natural recovery
and range expansion by comparison to areas where lynx have been
(mostly negative) attitudes toward large carnivores than might the artificially reintroduced (Bath et al., 2008; Kaltenborn, Bjerke,
general public (Bjerke et al., 2002; Ericsson & Heberlein, 2003; Nyahongo, & Williams, 2006; Lüchtrath & Schraml, 2015; Røskaft et al.,
Liukkonen et al., 2009; Kovařík et al., 2014). Considering small number 2003; Teel & Manfredo, 2010). Our findings did not support such an
of attacks caused by lynx on livestock per year (Kovařík et al., 2014; idea: we did not detect the hunters’ attitude being more positive in
Fig. S1) and existing compensation systems in many Central European areas with natural recolonization (E Moravia) compared to areas where
countries, hunters represent the key stakeholder group related to lynx lynx had been deliberately reintroduced (SW Bohemia), nor did atti-
conservation in this region. tudes vary between areas with higher (SW Bohemia) or lower lynx
The results from our survey could be slightly biased due to what population abundance (all other hunting areas).
might seem to be a relatively small response rate among hunters (al- Somewhat against our own expectations, the response of younger
though we reiterate that this will inevitably be underestimated due to people, exposed specifically to relevant education was not substantially
method of calculation, see Material and methods). However, the more positive than that of hunters. A half of the students at secondary
number of questionnaires returned is comparable to other similar stu- schools and a third of students at forestry faculties believed that lynxes
dies within Europe and the response rate was similar in surveys of both threaten roe deer stocks, even though red and roe deer generally occur
2001 and 2015. The main finding of this study is that the attitude of at relatively high abundance in the Czech Republic and caused ex-
Czech hunters toward lynx is predominantly negative regardless of the tensive damage in forest regeneration (Kamler, Homolka, Barančeková,
area, lynx population density or whether lynx populations arise from & Krojerová-Prokešová, 2010); 10% of university students are generally
natural range expansion or artificial re-introduction. It is clear, more- opposed to the re-establishment of lynx in the countryside.
over, that attitudes have become even more negative over the last 15 As with attitudes towards hunting in general (e.g. Floyd, Bankston,
years. Only approximately 20% of hunters perceived positive effects of & Burgesion, 1986; Stokke, 2004), a probable explanation for the per-
lynx on wildlife; the majority still believe that lynx have a negative sisting negative attitude of the students surveyed may derive from close
effect on wildlife, and 27% commented that they would prefer that no personal connection with family members or others who hunt; negative
lynxes were present in the environment. attitude to large carnivores seems to be deeply rooted and probably
This attitude strongly differs from that of general the public. A obtained in childhood (personal communication with students).
survey of the general public in the Czech Republic showed strong public

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4.2. Attitude towards legal hunting of lynx Poaching seems to be the most likely mortality factor preventing po-
pulation growth: our findings showed that hunters as well as forestry
However positive the attitude towards large carnivores by con- students are aware of illegal hunting being undertaken and a number of
servationists, conservation bodies, and general public, such a positive hunters admitted that they themselves had killed lynx on one or more
perspective is clearly not shared by key stakeholder groups such as occasions in all hunting regions.
hunters and livestock farmers. This difference in attitude is under- Firm estimates of the rate of illegal hunting are sparse in Europe.
standably due to differing socio-economic interests of different interest However, poaching is mentioned as the most common cause of lynx
groups (Lüchtrath & Schraml, 2015). mortality in Switzerland (Schmidt-Posthaus, Breitenmoser-Würsten,
Many hunters also have little confidence in the accuracy of scientific Posthaus, Bacciarini, & Breitenmoser, 2002), Poland (Jedrzejewski
estimates of population size. There is an enormous discrepancy in views et al., 1996), Croatia (Sindičić et al., 2016), and was admitted by
of the overall population size of lynx in the Czech Republic between hunters even in areas with really low lynx population density (e.g.
hunters’ estimates based on annual counts from individual hunting Macedonia; Lescureux et al., 2011). Andrén et al. (2006) estimated
grounds (Hunting Statistics of the Czech Republic 2000–2015; Fig. 6) poaching in Scandinavia to account for between 32 and 74% of total
and estimates offered from more objective scientific analyses. Hunters’ mortality in the Scandinavian lynx population.
estimates are more than three times higher than those obtained by One common method of quantifying causes of mortality in popu-
scientific monitoring based on camera trapping and noninvasive genetic lations of large carnivores is by assessing the fate of animals fitted with
sampling (Bartošová, 2004; Hlaváč et al., 2014; Krojerová et al. 2014; radio-transmitters (although this in itself makes the assumption that the
Poledníková et al., 2015; Heurich et al., 2018). Inevitably however, rate of mortality of animals fitted with such collars are unaffected by
hunters tend to believe in their own estimates and as a result, they may the collar, which may not be the case). However, when a radio-collared
be convinced that the lynx is overpopulated (stated as a cause of kill in animal is poached, there is a high probability that the poacher promptly
14% of surveys, Table S2), need to be managed by hunting and, since destroys the transmitter and hides the carcass, leaving the researcher
this is not currently possible within the law, they have resorted to with a lost signal without known cause (Goodrich et al., 2008) and thus
shooting lynx illegally. provides an underestimate of hunting mortality.
This perception of high population numbers underpins the hunters’ Another approach is to calculate the poaching rate through the
strong support for legal hunting of lynx (Fig. 3). Permitting controlled population growth modelling approach as unexplained mortality de-
hunting of predators may indeed improve attitude of hunters toward tected within the model (Heurich et al., 2018; Liberg et al., 2012).
these animals and make them a valuable game species (Linnell, Based on population modelling, we calculate that an annual mortality
Swenson, & Andersen, 2001; Heberlein & Ericsson, 2008; Treves, 2009). in the region of 35% would be required if populations within the Czech
Management of lynx by hunting was considered as one of the most Republic have remained stable and shown zero growth. Natural mor-
important methods for improving human-lynx coexistence in Finland tality is known to be 2–5 % (Andrén et al., 2006; own data) while ac-
by all stakeholder groups except for conservationists themselves (who cidental death through involvement in collisions with vehicles may
promote education as the most important method) (Liukkonen et al., form similarly up to 5% of the total mortality (Wölfl et al., 2001). On
2009). This approach already works for the bear in Finland (Mykrä the presumption that a 10% loss results from natural mortality and
et al., 2017). accidental death in this way, we calculate that at least 25% of adults
However, other studies, especially those about more controversial might be killed illegally each year. An even higher estimate of the rate
species such as wolves, did not detect a change of attitude in positive of illegal killing would if fecundity rates (higher number of kittens per
direction even when some degree of legal hunting was permitted litter) or reproductive rates (higher proportion of reproducing females)
(Andrén et al., 2006; Hogberg et al., 2016). Moreover, it was detected are in practice higher than those we have assumed in our simulations.
that harvesting may negatively influence the ecological role of these Similar estimates of rates of illegal killing (of between 18–20 %) were
apex predators (Ordiz, Bischof, & Swenson, 2013). At present, as noted recently reported from the study of Heurich et al. (2018).
in our Introduction, the lynx is strictly protected by international and
national laws in all Central European countries. Further, the dis- 5. Conclusions
continuous distribution coupled with the small current population size
of lynx populations in Central Europe render it impractical for man- Illegal hunting seems to be the most important factor influencing
agement strategies to encompass legal hunting within the immediate, or rates of population growth of all European lynx populations and can
foreseeable, future. greatly increase the probability of extinction, especially of small-sized
or founder populations (Nowell & Jackson, 1996; von Arx et al., 2004;
4.3. Illegal hunting of lynx Sindičić et al., 2016; Heurich et al., 2018). Illegal hunting of lynx occurs
in all European countries where lynx occur regardless of living stan-
Because of their low densities combined with their slow rates of dards or education (Andrén et al., 2006; Dressel et al., 2015). It is ap-
population growth, top predators are particularly vulnerable to parent that the ultimate motivation is a lack of acceptance of the
poaching (Liberg et al., 2012). Almost all large carnivore species have human-lynx coexistence in the modern landscape by key stakeholders
endured a long history of human persecution and have been eradicated (Andrén et al., 2006). Therefore, the main task to ensure long-term
from substantial parts of their historical ranges. Even though they are survival of lynx in Europe is to increase tolerance towards the lynx
now protected in many areas, poaching still represents a widespread thereby reducing poaching impact.
problem for their conservation (Woodroffe & Ginsberg, 2000; von Arx, Because a substantial part of poaching is often unobserved,
Breitenmoser-Würsten, Zimmermann, & Breitenmoser, 2004; Chapron poaching may be an even larger problem in wildlife conservation than
et al., 2014; Liberg et al., 2012; Gangaas et al., 2013; Heurich et al., has been previously assumed. By contrast to farmers who may receive
2018). It follows that dealing with illegal hunting often emerges as a financial compensations for their loss of livestock (even though com-
necessary condition for the restoration, conservation and sustainable pensations received may be less than the real damage), hunters get
management of large carnivore populations (Heurich et al., 2018; nothing for their loss of wild game, whether this loss is perceived or
Liberg et al., 2012). actual (in reality, hunters have never had to reduce hunting quotas due
Objective estimates of the size of the lynx population in the Czech to lynx predation in any hunting grounds in the Czech Republic).
Republic from scientific analyses indicate oscillation of population Further, many of them still consider hunting of large carnivores as a
numbers around a stable value instead of the increase in population part of their entitlement and also as an opportunity to obtain a valuable
numbers which might have been expected over the last 15 years. trophy.

35
J. Červený et al. Journal for Nature Conservation 47 (2019) 28–37

Even though the current rate of illegal hunting in the Czech Breitenmoser, U., Breitenmoser-Würsten, C., Okarma, H., Kaphegyi, T., Kaphegyi-
Republic seems to keep the lynx population at the same level rather Wallmann, U., & Müller, U. M. (2000). Action plan for the conservation of the
Eurasian lynx (Lynx lynx) in Europe. Council of Europe, Nature and Environment, 112,
than causing actual decline at a national level, poaching is un- 1–69.
predictable and beyond any control. Based on experience of recent Breitenmoser, U., Ryser, A., Molinari-Jobin, A., Zimmermann, F., Haller, H., Molinari, P.,
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The study was financially supported by EEA grants (project MGSII- young adults’ social experience on their attitudes toward hunting and hunters.
40) and by Institutional Research Plan (RVO: 68081766). Journal of Sport Behavior, 9, 116–130.
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Acknowledgements Ecology, 83, 107–115.
Gangaas, K. E., Kaltenborn, B. P., & Andreassen, H. P. (2013). Geo-spatial aspects of
We would like to thank Elisa Belotti, Luděk Bufka, Miloš Ježek, Jiří acceptance of illegal hunting of large carnivores in Scandinavia. PloS One, 8(7),
e68849.
Kamler, and Petr Konupka for their help with surveys distribution and/ Goodrich, J. M., Kerley, L. L., Smirnov, E. N., Miquelle, D. G., McDonald, L., Quigley, H.
or other reliable data and comments useful during manuscript pre- B., et al. (2008). Survival rates and causes of mortality of Amur tigers on and near the
paration. Also we would like to thank to Rory Putman for revision of Sikhote-Alin Biosphere Zapovednik. Journal of Zoology, 276, 323–329.
Heberlein, T. A., & Ericsson, G. (2008). Public attitudes and the future of wolves Canis
English and all valuable comments and suggestions to previous versions
lupus in Sweden. Wildlife Biology, 14, 391–394.
of the manuscript. Further, we thank editor and two anonymous re- Hell, P., Slamečka, J., & Gašparík, J. (2004). Rys a divá mačka v slovenských Karpatoch a vo
viewers for their valuable suggestions that contributed to the im- svete. Bratislava: Slovakia: PaRPress (in Slovak).
Heurich, M., Schultze-Naumburg, J., Piacenza, N., Magg, N., Červený, J., Engleder, T.,
provement of final version of the manuscript.
et al. (2018). Illegal hunting as a major driver of the source-sink dynamics of a re-
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