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European Journal of Wildlife Research (2022) 68:9
https://doi.org/10.1007/s10344-021-01556-5
ORIGINAL ARTICLE
Conflicts betweenlarge carnivores andlocal pastoralists
aroundNiokolo Koba National Park, Senegal
MalléGueye1,2,3· DorienVanCauteren2· LisaMengual2· RaoulPellaton2· HerwigLeirs2· LauraD.Bertola4,5·
HansdeIongh2,6
Received: 14 August 2020 / Revised: 4 December 2021 / Accepted: 20 December 2021
© The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature 2022
Abstract
In Niokolo Koba National Park (NKNP), in the South East of Senegal, we studied the current spotted hyena (Crocuta crocuta)
population size, density and distribution and assessed human-large carnivores’ conflicts. We implemented 26 calling stations
operations, installed 87 camera traps and obtained data from tourists, local guides and rangers. In addition, a questionnaire
survey was carried out in 31 villages around the park. Our results showed an estimated density from 1.77 to 3.53 spotted
hyenas/100 km2 depending on used response rates (50–100%) with a minimum population size between 144 and 288 indi-
viduals. According to the query survey, the spotted hyena is responsible for the majority of livestock predation around the
park (87% of all attacks on livestock), followed by African wild dog (6%), lion (4%) and leopard (3%). A total of 63.1% of
all attacks were on cattle, while sheep and goats represent the remaining 36.9%. Spotted hyena and lion mainly predated on
livestock at night (93.5% and 71.4% of all attacks, respectively) in contrast to predation by African wild dog and leopard,
for which the majority of the attacks occurred during the day (63.9% and 62.5%, respectively). Furthermore, there was more
livestock killed during the wet season (58% of all attacks) than during the dry season (42% of all attacks). The distribution of
conflicts across five sectors surrounding the park illustrate that the West sector is most impacted (49% of all attacks; districts
of Linkering and Medina Gounass) followed by the North West (32%; districts of Dialocoto and Missira). Insights into the
severity and dynamics of human-wildlife conflict will contribute towards finding effective mitigation measures, minimising
loss of livestock and reducing conflicts.
Keywords Human-wildlife conflicts· Large carnivores· Spotted hyena (Crocuta crocuta)· Senegal· West Africa· Niokolo
Koba National Park
Introduction
Large mammal populations are declining in many African
countries, especially in areas with high human density, lead-
ing to extensive habitat destruction and bush meat poach-
ing (Craigie etal. 2010; Inskip and Zimmermann 2009).
Large carnivores are especially vulnerable as they often suf-
fer prosecution when in close proximity to humans. They
are generally declining worldwide (Ripple etal. 2014), but
trends vary according to geography (Chapron etal. 2014)
and the severity of threats posed to humans (Gompper
etal. 2015). Many species therefore depend on conserva-
tion measures such as protected areas. Due to the often
small size of these protected areas, the balance between
large predators and their prey populations is delicate and
easily disturbed. When natural prey is depleted, e.g. as a
result of bush meat hunting, carnivores may increase their
This article is part of the Topical Collection on Road Ecology
Guest Editor: Marcello D’Amico
* Mallé Gueye
mallegueye@yahoo.fr
1 Niokolo Koba National Park, BP 37, Tambacounda, Senegal
2 Department ofBiology, University ofAntwerp, Campus Drie
Eiken, Universiteitsplein 1, B-2610Wilrijk, Belgium
3 Department ofHydroSciences andEnvironment, University
Iba Der Thiam ofThiès, site VCN, Thiès, Sénégal
4 City College ofNew York, 160 Convent Ave, NewYork,
NY10031, USA
5 Section forComputational andRNA Biology, Department
ofBiology, University ofCopenhagen, CopenhagenN2200,
Denmark
6 Institute ofEnvironmental Sciences Leiden, Einsteinweg 2,
Leiden, TheNetherlands
European Journal of Wildlife Research (2022) 68:9
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9 Page 2 of 11
home ranges or shift their activities outside protected areas,
resulting in conflicts with the surrounding livestock owners
(Inskip and Zimmermann 2009; Lozano etal. 2019; Kissui
etal. 2019; Hazzah etal. 2014).
Also, in most of Africa’s protected areas, large mam-
mal populations have shown declining trends over the past
decades, with the strongest decline in West Africa (Craigie
etal. 2010). In West Africa, large carnivores have greatly
declined in numbers, and have also become increasingly
rare outside protected areas (Bauer and de Iongh 2005;
Riggio etal. 2012; Henschel etal. 2014). For carnivores,
the carrying capacity of a protected area is determined by
prey availability and vegetation, and with habitat destruc-
tion and declining prey populations, predators will attempt
to find other food sources outside of the protected area in
order to survive (Lesilau2019). As a consequence, the
home ranges of large carnivores may increase with a short-
age of prey (Bauer and De Iongh 2005; Van Oijen and
Rougier2005; Tuqa etal. 2014; Lesilau etal. 2018).
In areas where humans and carnivores live in close
proximity, the carnivores’ diet often includes livestock
(Woodroffe and Frank 2005; Tuqa 2015; Tumenta 2012).
Human-carnivore conflicts inevitably occur in a landscape
where people, wildlife and carnivores are forced to live
together (Inskip and Zimmermann 2009; Lozano etal.
2019). As a response to this conflict, people may resort
to retaliatory killing of these carnivores. The active per-
secution of carnivores by humans, as a result of conflict,
is the main reasons for population declines of both large
carnivores and their prey (Treves and Karanth 2003). The
conservation of large carnivores depends largely on the
desires and decisions of the local communities, as they
bear the costs of human-carnivore conflicts but could
also potentially participate in benefits from tourism and
employment (Kolipaka 2018). Pastoralists often make an
effort to reduce losses to large carnivores, mostly consist-
ing of keeping livestock in enclosures at night and herding
during pasture. However, these measures appear not to be
sufficiently effective (Yirga 2013).
There is a large number of scientific publications on
human-carnivore conflicts with large carnivores such as
lion, leopard and cheetah in different regions in Africa
(Inskip and Zimmermann 2009). However, there have been
few studies on human-carnivore conflicts with spotted
hyena (Crocuta crocuta, Erxleben 1777) in West Africa
(Di Silvestre etal. 2000). This was one of the main rea-
sons to implement a study on spotted hyena conflicts in
and around Niokolo Koba National Park (NKNP), Senegal.
Much of the background information comes from other
regions in Africa.
Spotted hyenas are largely nocturnal, with individuals
often moving long distances when active (Kruuk 1972;
Mills and Hofer 1998). Spotted hyenas are opportunistic
carnivores, both killing and scavenging on a wide range of
prey species (Cooper 1999).
The population of spotted hyenas in Africa ranges from
27,000 to 47,000 mature individuals (Mills and Hofer 1998).
This makes them probably the most abundant large carni-
vore of the continent, despite being consistently persecuted
(Yirga 2013). For West Africa, no population estimates are
known, yet for NKNP, Senegal, the local population size
of the spotted hyena was estimated at between 200 and 500
individuals (Sillero-Zubiri etal. 1997), with no more recent
estimates available. Most spotted hyena populations in pro-
tected areas of southern Africa are stable, but in protected
areas of eastern and western Africa, they are declining due
to incidental snaring and poisoning (Mills and Hofer 1998;
Craigie etal. 2010). Diseases, habitat loss and persecution
by humans make spotted hyenas dependent on conserva-
tion areas, especially in West Africa (Mills and Hofer 1998;
Bohm and Höner 2015; Sogbohossou etal. 2018; Green and
Holekamp 2019; Kirsten etal. 2017). Various mitigation
measures are known to reduce the chances of livestock dep-
redation. In general, livestock that is closely herded by day
and kept in bomas at night with watch dogs and high human
activity are less likely to be attacked (Lesilau etal. 2018).
Our study aimed at a population assessment of spotted
hyena in NKNP and an analysis of associated conflicts with
livestock owners. In order to achieve these aims, the follow-
ing research questions were set:
a) What is the current spotted hyena population size, den-
sity and distribution in the NKNP?
b) What are the trends of human-hyena conflicts in time
and space in comparison with other large carnivores?
c) Which independent factors influence human-hyena con-
flicts?
Materials andmethods
Study area
NKNP is located in the south east of Senegal (Fig.1) and
was established as a national park in 1954. It is currently
on the list of UNESCO World Heritage in Danger. With its
9130 km2 (8130 km2 being the core area), it is one of the
largest national park of West Africa (Madsen etal. 1996).
The park is situated in the west Sudan savanna zone (Magin
2004) and contains woodland savanna and seasonal wetlands
(Madsen etal. 1996).
NKNP and the adjacent protected areas in Guinea are
one of the last remaining protected savanna landscapes in
West Africa. It is the habitat for about 330 bird species and
80 mammals species, including monkeys, warthog (Phac-
ocheorus aethiops), hippos (Hippopotamus amphibius)
European Journal of Wildlife Research (2022) 68:9
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and several large predators: West African lions, African
leopard, African wild dog and spotted hyena (Kane 2014).
It further hosts more than 1500 species of flowering plants
(Adam 1971; Schneider and Sambou 1982).
The direct surrounding of the park is inhabited by people
from different ethnic groups. Some people herd their livestock
in enclosures while others let them wander freely. The villages
are located outside the park boundaries but some of them are
Fig. 1 Map of Niokolo Koba National Park. Localities of camera traps and calling stations are indicated. The map also includes the buffer zone
surrounding the park, the villages selected for the survey and the five sectors used for further analysis
European Journal of Wildlife Research (2022) 68:9
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located close to the buffer zone, making encounters with wild-
life relatively common. The buffer zone is a stretch of 1000m
width surrounding the entire park, in which there are no human
settlements, but where human activities are allowed.
Average annual rainfall ranges from 900 to 1200mm
from May to October and temperatures range from 25 in
December to 33°C in May (DPNS 2000). That corresponds
to a tropical climate with dry season in winter (Aw) accord-
ing to the climatic classification of Köppen (Gueye 2014
thesis report). NKNP is provided with water by the Gambia
River and its tributaries, theNiokolo-Koba and Koulountou.
Data collection
Calling station survey
The population size of spotted hyena was estimated with a
calling stations survey conducted from 9 to 19 April 2018.
The operation covered 26 calling stations’ representative for
the study area (Fig.1). We used an adapted method accord-
ing to Ogutu and Dublin (1998), with four times 15-min
African buffalo distress calls and five-minute silence in
between each sequence, transmitted through a megaphone
(Flextone FLX1000) positioned on the roof of a vehicle.
The response distance was considered to be 3km, based on
response distances reported in literature (Sogbohossou and
Téhou2007; Omoya etal. 2014), thus representing an area
of 28.27 km2 per calling station. The interval between two
neighbouring calling stations was fixed at 7km to avoid
overlap and double counting. According to Bauer (2007),
density calculations should ideally be based on local cali-
brations with an empirical assessment of response distance
and response rate. That is, one should find large carnivore
individuals first, then call from 5km and progressively
come towards them and assess the distance at which they
react (response distance), as well as the percentage that
reacts at all (response rate). This was practically impossi-
ble, which is why we used figures from literature instead.
Ogutu and Dublin (1998) found a response distance of
2.5km and a response rate of 26%. Mills etal. (2001) found
a response distance of 3.2km and an individual response
rate of between 43 and 77% for hyenas. Calling station sur-
veys in Bouba Ndjida National Park, Cameroon, estimated
a response distance of 2.5km and assumed a 75% response
rate in similar habitat conditions as in NKNP (Bauer 2007).
We used a conservative estimate of 3km for the response
distance and calculated the densities and population size for
a range of response rates, including 50%, 75% and 100%.
Camera traps
Data collected with 87 camera traps (Fig.1) used in the
ecological monitoring programme between 2016 and 2019
was used to create a distribution map of spotted hyena in the
park. Furthermore, data collected from tourists, guides and
rangers on the observations of the species were included.
We designed a 4-km2 grid pattern covering NKNP and
then divided the park into 15 zones. In each zone, 3 to 20
grid cells were sampled by placing a camera inside for 1 to
3months. The sampled grids were selected based on the
presence of animal tracks, vehicle accessibility and water
point proximity. This approach is expected to give a more
reliable estimate, since placement on dirt roads or large
game tracks has been shown to influence capture rates of
large carnivores (Trolle and Kery 2005; Weckel etal. 2006).
Questionnaires
An adapted semi-structured questionnaire was carried out.
Villages for the questionnaire survey were selected based on
their proximity to the border of the park, distribution over the
five different sectors of the park (west, north east, north west,
south and south east) and accessibility by car. We selected
villages within 10km from the park boundary and population
data were collected at the local community department. In
each village, we semi randomly selected a minimum of 5%
of the household heads for interviews (Grosh and Glewwe
2000; Kulpa and Szarata 2016). We assumed that this sample
size was representative for the whole village. If the house-
hold head was not present, we interviewed another senior
member of the household. In total, 204 respondents in 31
different villages were interviewed between March and June
of 2018. Figure1 provides an overview of the villages that
were visited, showing the distribution around the border of
the park. In the questionnaire survey, respondents reported
their livestock losses and the large carnivores believed to be
responsible for each attack. The studied livestock included
cattle, sheep and goats (shoats). Pictures of animals were
shown to facilitate identification of predators and additional
background information was provided. When an attack on
livestock occurred, the respondents confirmed that they
could identify the predator by either direct sighting or by
tracksin the ground and bite marks on the killed livestock.
In all cases, respondents identified the correct carnivores,
as this was checked by the researchers. The interviews were
done during the dry season (March–June), yet questions of
the interview referred to all seasons.
Data analysis andstatistics
Obtained data was analysed with MS Excel software. Spotted
hyena density was estimated by dividing up total number of
individuals observed by the sum of 3km circles surrounding
calling stations. The obtained value is related to the response
rate (50, 75 and 100%). Regarding the population size, it was
estimated by extrapolation of the density on the core area
European Journal of Wildlife Research (2022) 68:9
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of NKNP. In order to estimate the total reported livestock
loss for the whole village, we extrapolated the losses of the
sample of households to the total number of households in
each village. The total fraction of livestock lost was based on
the estimate of livestockdepredationby respondents during
the past year, in relation to the number of cattle and shoats
they own. These numbers included all reported attacks on
livestock, both inside and outside the park.
The following independent factors were included: (1)
season (wet and dry season defined as June to December for
the wet season and January to May for the dry season), (2)
distance to park boundary in km, (3) time of the attack as
during nighttime (18–06h) or daytime (06–18h).
R software was used to assess the VIF (variance inflation
factor) to determine which variables were correlated and which
variables may be included in the model. Linear and multiple
regressions were carried out in order to assess how the differ-
ent variables (number of cattle, number of shoats, distance to
the park, population) impacted the number of total attacks.
The linear and multiple regression analysis were per-
formed after checking the mainassumptions for these analy-
ses (independence of observations, normality and linearity,
see Suppl. Fig.1 for the test of collinearity).
In addition, we did a principal component analysis (PCA)
based on several variables (sectors, number of attacks on
cattle, on shoats, total attacks, ethnic groups, villages’ geo-
graphic coordinates, distance to the park boundaries). This
method of analysis aims to reduce the number of variables
in the data sets, to simplify the observations while retaining
as much information as possible. An ascending hierarchi-
cal classification was also performed in order to cluster vil-
lages according to their respective predation intensity. The
Shapiro–Wilk test was used to indicate the variables whose
modalities best separated the villages on the plane.
Results
Spotted hyena population size, density
anddistribution
The total area of hyena habitat covered by the 26 calling sta-
tions was 735.11 km2 (9.04% of the core area of NKNP, see
Fig.1), based on a response distance of 3km. The average
hyena density in this area was calculated as 3.53, 2.36 and
1.77 spotted hyena per 100 km2, depending respectively on
the used responses rate 50%, 75% and 100%. This suggests
that the minimum population size of the spotted hyena is
within the range of 144–288 individuals in the core area of
NKNP.
The database analysis of fauna observations combined
with the camera traps survey showed that the spotted hyena
is widely distributed in NKNP. It is present in all zones and
both inside the park and in the border area of the protected
area. However, individuals are more frequently observed in
the South and South East sectors.
Livestock depredation andlarge carnivores involved
According to the responses to our survey, the predation
intensity seems to have increased from two cases in 2013
to 210 cases in 2017 (Supplementary Table1). Unfortu-
nately, there have not been previous surveys, and therefore,
a direct comparison to assess livestock depredation through
time was not possible. We acknowledge that older estimates
may be inaccurate; however, we consider the differences in
reported livestock depredation large enough to conclude
that an increase is very likely. The large carnivores involved
in attacks on cattle and shoats were spotted hyena (87% of
attacks), African wild dog (6%), West African lion (4%)
and African leopard (3%). Furthermore, in 2016, no leopard
attacks were recorded. A proportion of 63.1% of all attacks
concerned cattle, while shoats were attacked in 36.9% of
cases. Thus, the number of predations increased from 2013
to 2017 and spotted hyena was reported to be responsible
for most livestock mortalities recorded on cows and shoats
around the park (n = 184).
Based on the answers of the respondents, people believed
that cattle were most often killed by spotted hyena (81%)
followed by African wild dog (8%), lion (7%) and leopard
(4%). Similarly, shoats were believed to be attacked primar-
ily by spotted hyena (93%), while African wild dog (5%)
and leopard (2%) were only marginally responsible. There
was no documentation of lions attacking shoats around the
park (Fig.2).
Fig. 2 Fraction of reported
livestock killed and contribu-
tion of different large carni-
vores on livestock depredation
events between 2013 and 2017
(n = 184) around Niokolo Koba
National Park
81%
7%
4%
8%
(A) Cattle lost
Spotted hyenaLion Leopard African wild dog
93%
2% 5%
(B) Shoats lost
Spotted hyena Leopard African wild dog
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Livestock depredation tolarge carnivores intime
andspace
According to the respondents, predation intensity of large
carnivores around the park seemingly varied in time and
season. Attacks were frequently observed during the night
between 00:00 and 06:00 (53%) and 18:00 and 00:00 (23%)
(Fig.3). Spotted hyena (93.5%) and lion (71.4%) were
reported to mainly predate livestock at night while predation
by African wild dog and leopard was typically most intense
during the day (63.9 and 62.5%, respectively). There was
more livestock killed during wet season from May to Octo-
ber (58%) than during the dry season (42%) from November
to April.
Geographical distribution andintensity
ofpredation
Following the results of the questionnaire, during 2013 to
2017, the number of large carnivore predation events was
most important in the West sector of the park (49%), fol-
lowed by the North West (32%) and the South (8%). The
South East and North East sectors recorded respectively 6%
and 5% of livestock predation. These results are represented
in Fig.4 showing the geographical distribution of the preda-
tion and its intensity for each studied village around the park.
Using VIF, the variables Total_Attacks and Attacks_on_
Cattle (R2 = 0.92) and Total_Attacks and Attacks_on_Shoats
(R2 = 0.84) were reported to be strongly correlated, as were
the Attacks_on_Cattle and Total_Cattle (R2 = 0.77). Correla-
tions between other variables were relatively low (R2 < 0.7)
(Suppl. Fig.1).
We found a significant relationship between the total
number of attacks and the number of cattle(p-value < 0.001),
with a 0.0017 unit (± 0.00036) increase in reported attacks
for every unit increase in cattle (Fig. 5, left panel). In
06-12h
12%
12-18h
12%
18-00h
23%
00-06h
53%
06-12h 12-18h 18-00h 00-06h
Fig. 3 Time of large carnivores attacks around Niokolo Koba
National Park, based on data from 2013 to 2017 (n = 184)
Fig. 4 Location of camera trap recordings of spotted hyena and reported incidences and magnitude of livestock depredation between 2013 and
2017 around Niokolo Koba National Park
European Journal of Wildlife Research (2022) 68:9
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addition, we found a similar effect between total number
of attacks and the number of shoats(p-value < 0.01), with
a 0.0026 unit (± 0.00073) increase in attacks for every unit
increase in shoats (Fig.5, right panel). There was no signifi-
cant effect of distance to the park or population (i.e. village
size).
The critical probability of the Shapiro–Wilk test indicated
that the variable sectors or subdivisions of the park was the
only qualitative variable to illustrate the distances between
villages yet was not significant (p = 0.1365368). A PCA was
performed using the numbers of attacks on cattle, shoats and
total attacks (Suppl. Fig.2). Villages were coloured accord-
ing to their sector affiliation. The first dimension of the PCA
opposes villages such as Kalifourou and Wadiatoulaye to
villages like Gamon, Kouar Saal, Mithiou, Nieminike and
Mansadala.
The questionnaire survey distinguished four main eth-
nic groups, namely Peul, Bassari, Malinké and Tendanké,
with some villages representing a mix of ethnic groups.
According to this study, the Peul ethnic group, being tra-
ditional breeders, is most strongly represented around the
park. Figure6 shows that livestock attacks were predomi-
nant in the West, North West and South East of NKNP,
where respectively Peul, Tendake and Malinke ethnic
groups were most impacted. The South East, inhabited
by Bassaris, recorded the least attacks compared to the
other sectors.
The classification carried out on the intensity of preda-
tion in villages around the park shows five classes (Suppl.
Fig.3).
Class 1 is made up of villages such as Gamon, Mansadala,
Kouar Saal, Mithiou, Woundiou, Nieminike and Islam.
This group is characterised by low values for the variables
“total number of predation”, “number of cattle killed” and
“number of shoats killed” (from the most extreme to the
least extreme);
Fig. 5 Linear regression of total reported attacks and number of cattle (left) and number of shoats (right)
Fig. 6 Number of livestock
attacks per village for data
collected 2013–2017 (n = 184).
Data are represented per ethnic
group, with colours indicating
how these ethnic groups are
distributed over the five sectors
surrounding Niokolo Koba
National Park
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Class 2 is made up of villages such as Kouar and Badon.
This group is characterised by variables whose values do
not differ significantly from the mean;
Class 3 is made up of villages such as Belly, Akkan and
Samaye. This group is characterised by high values for the
number of cattle killed;
Class 4 is made up of villages such as Dienoudiala and
Medina couta. This group is characterised by high values
for the number of shoats killed;
Class 5 is made up of villages such as Dar Salam, Kali-
fourou and Wadiatoulaye. This group is characterised by
high values for the variables total number of predations,
number of shoats killed and number of cows killed (from
the most extreme to the least extreme).
Discussion
Spotted hyena population size, density
anddistribution
The minimum population size of spotted hyena in the
park was estimated at 144–288 individuals, based on three
response rates (50%, 75% and 100%) which is lower com-
pared to the findings of Sillero-Zubiri etal. (1997), who
estimated the population between 200 and 500 individu-
als in NKNP. It shows that numbers have become unsta-
ble and decreased over the last 25years. We acknowl-
edge that a direct comparison of spotted hyena densities
in NKNP with estimates from others national parks may
contain some bias if the density estimates were not based
on response distance (3km) and response rate (50%, 75%
and 100%) used in this study. These data therefore only
give an indication of differences in density.
However, in other West African national parks such as
the WAP Complex (on the border of Benin, Burkina Faso
and Niger), the estimated population of spotted hyena is
1119 individuals (Arly-Pendjari, n = 723 individuals; W
National Park, n = 396 individuals) (Bouché etal. 2014).
Henschel etal. (2014) suggested that the WAP shelters an
important population of spotted hyenas in West Africa.
In addition, the Bénoué Complex in Cameroon hosts a
population of about 1376 individuals (range 1352–1400)
(Bénoué, Bouba Ndjida (87–618 spotted hyena for the
southern sector, Faro National Park and 32 hunting zones))
(Bauer etal.2016). In Dinder National Park, Sudan, the
population size is estimated between 36 and 89 individuals
(Mohammed etal. 2019).
Regarding the population density based on the calling
station survey, our estimations of 3.53 spotted hyenas/100
km2 show that NKNP records lower spotted hyena density
than the Bénoué Complex with 5.69 individuals/100 km2
(Bauer etal. 2016), but higher than the Dinder National
Park (1.62–1.98 individual/100km2) (Mohammed etal.
2019) based on different density estimation methods.
However, with other spotted hyena estimating den-
sity methods, the WAP Complex hosts about 4.3 indi-
viduals/100 km2 (Bouché etal. 2014), the Maasai-Mara
Serengeti ecosystem (60–80 individuals/100km2) and the
Kruger National Park 7–20 individuals/100 km2 (Hunter
and Barrett 2011). Some further characteristics of spotted
hyena population across Africa are summarised in Sup-
plementary Table2.
Our results show that spotted hyenas have a wide distri-
bution within NKNP. Spotted hyenas are present in differ-
ent parts of the park, particularly in the centre and in the
buffer zone (see Fig.3). This pattern could be explained by
the availability of wildlife prey in the core area and live-
stock prey in the buffer zone. Rabeil etal. (2018) showed
in the most recent wildlife census in NKNP that wildlife
densities in the park are lower in areas where anthropo-
genic activities (pasture) are high, suggesting that human
disturbance has a direct negative impact on prey densities.
In addition, the plasticity of the diet of the spotted hyena
may explain its wide distribution. Spotted hyenas are oppor-
tunistic and their high densities can be partly explained by the
presence of lions and lion kills. When prey is scarce, the spot-
ted hyena has the ability to travel more than 10km in a night
or day to search for prey (Yirga 2013). In Maasai Mara, hyenas
travel 12.4km on average per night (Kolowski etal. 2007).
Livestock depredation andlarge carnivores involved
In our study, the trend of the number of attacks by large
carnivores on cattle, sheep and goats was increasing around
the park from 2013 to 2017. Spotted hyena, African wild
dog, lion and leopard were all involved in livestock preda-
tion. However, the spotted hyena was considered the pre-
dominant predator of livestock. The dominant role of spotted
hyena can be explained by the high behavioural plasticity of
spotted hyena (Boydston 2003). According to some studies,
spotted hyenas have a reputation of killing and scavenging
domestic livestock, mostly cattle, sheep and goats (Mills and
Hofer 1998; Yirga etal. 2011).
In terms of conflicts, the spotted hyena is reported to be
the main responsible for livestock depredation around the
park (87% of all attacks on livestock), although other car-
nivores were reported to be involved in livestock depreda-
tion as well. The size of the livestock killed by these large
carnivores corresponds to the typical size of natural prey
(Patterson etal. 2004 in Bodendorfer etal. 2006; Bauer etal.
2008 in Sogbohossou etal. 2011).
The African lion is an opportunistic hunter, but it gen-
erally prefers larger herbivores weighing between 60 and
550kg body mass (Hayward and Kerley 2005). Spotted hye-
nas are generally nocturnal and, in our study, most attacks
European Journal of Wildlife Research (2022) 68:9
1 3
Page 9 of 11 9
happened during the night. It is known that spotted hyenas
kill their own prey in 60 to 95% of cases (Kruuk 1972). Afri-
can wild dog attacks on cattle can be justified by the behav-
iour of this species operating in groups, so that they are able
to kill larger prey. It also frequently hunts smaller prey, espe-
cially when large species are absent or rare, such as outside
protected areas. It rarely kills small cattle (sheep, goats, etc.)
if wild prey is available (Hunter and Barrett 2011).
Livestock depredation tolarge carnivores intime
andspace
Spotted hyena and lion mainly attacked livestock at night
(93.5% and 71.4%, respectively) in contrast to African wild
dog (36.1%) and African leopard (37.5%). The attacks of
African wild dog and leopard are consequently most intense
at day, possibly in order to avoid possible competitions with
lion and spotted hyena. Overall however, our results show
that the intensity of predation is higher during the night. The
spotted hyena is known to be most active during the night
(Kruuk 1972; Mills and Hofer 1998) as well as the lion,
which is mainly a crepuscular to nocturnal animal (Hunter
and Barrett 2011). There was more livestock killed during
the wet season (58%) than during the dry season (42%). A
possible explanation may be the fact that in the wet season,
the natural prey of large carnivores is no longer concen-
trated around water points but rather dispersed throughout
the park, given a good distribution of rainfall. Therefore,
large carnivores attack livestock to fill the temporal lack of
natural prey by moving in and out of the park, increasing
their attacks on livestock. Various studies indicated seasonal
variation in movements of spotted hyenas due to fluctuations
in prey abundance (Hofer & East 1993; Trinkel etal. 2004 in
Yirga etal. 2011; Tumenta 2012; Tuqa 2015; Lesilau2019).
In northwestern Zimbabwe, lion and hyena attacks on cattle
were more frequent in the wet season (Loveridge etal. 2017;
Croes etal. 2011; Bakker etal. 2020).
Geographical distribution andintensity
ofpredation
We found that the livestock predation by large carnivores is
present all around the park with different levels of impact. The
most impacted local communities tended to have larger live-
stock herds, reside closer to the park where they often allow
their livestock to graze on pastures inside the park. We recorded
attacks per village, but it does not mean that the livestock was
attacked in the village; it could also have been livestock from
the village that was taken for grazing inside the park.
The West sector followed by the North West is the most
strongly impacted by large carnivore attacks compared to
the South, the South East and the North East of the park.
This may be explained by the population composition around
NKNP. West and North West sectors are inhabited by a human
population composed essentially of the Peul ethnic who are
traditional cattle breeders. Indeed, in 2012, Linkering, Medina
Gounass and Dialocoto recorded respectively 36,000, 22,600
and 29,000 individuals of livestock (cattle, sheep and goats)
based on data from the Veterinary Services of Tambacounda
and Kolda in 2014. Our results indicate a positive correla-
tion between number of cattle and number of shoats, and the
number of attacks (Suppl. Fig.1), and the highest number
of attacks/village has been recorded in villages which were
mainly inhabitated by the Peul ethnic group (Fig.6).
In the West sector, the villages Kalifourou and Wadiatou-
laye are more impacted by large carnivore predation, both on
cattle and shoats, than Gamon, Kouar Saal, Mithiou, Niemi-
nike and Mansadala. Indeed, the first two mentioned villages
belong to the Fouladou area which is a region principally
inhabited by Peul.
Specifically, the low numbers of cattle and shoats killed
in Gamon, Mansadala, Kouar Saal, Mithiou, Woundiou,
Niminike and Islam could possibly be due to the protec-
tive measures taken by breeders to keep livestock in wooden
enclosures and shepherds accompanying them while pas-
turing. High predation values in the villages of Dar Salam,
Kalifourou and Wadiatoulaye can be explained by the size
of livestock herds, the proximity to the park and the lack of
natural prey in those sectors. Furthermore, villages char-
acterised by high numbers of cows or shoats killed can be
explained by the large availability of livestock and a lack of
protective measures.
Conclusion
We conclude that depredation of livestock by spotted hyena
in and around NKNP has by far the largest impact on live-
stock owners living around the park. Predation by lion, leop-
ard and African wild dog are negligible compared to the
number of attacks by spotted hyena. Livestock was mainly
predated at night and attacks were higher during the rainy
season. The most impacted local communities tended to
have larger livestock herds, reside closer to the park where
they often pasture their livestock. The results of this study
may be used for the conservation of wildlife in the area and
for the protection of local populations’ livestock.
Supplementary Information The online version contains supplemen-
tary material available at https:// doi. org/ 10. 1007/ s10344- 021- 01556-5.
Acknowledgements We acknowledge Dr Fatou Nene Diop, for sta-
tistical analysis, and Lt Maniang Mamadoune Diop and Lt Papa Mor
FAYE for collecting some of the data. We also thank the National Park
Department and all rangers who contributed to the study.
European Journal of Wildlife Research (2022) 68:9
1 3
9 Page 10 of 11
Author contribution All the authors have contributed to data collecting
or analysis and improving the manuscript.
Funding The research was supported by the Kaplan Panthera Grant
and the Leo Foundation.
Availability of data and material The main data were collected in the
field by Dorien Van Cauteren, Lisa Mengual, Raoul Pellaton and Mallé
Gueye. They are available with the first author.
Declarations
Ethics approval Not applicable.
Consent to participate All authors consent to participate in the writing
of this article entitled “Conflicts between large carnivores and local
pastoralists around Niokolo Koba National Park, Senegal”.
Consent for publication All authors consent to publish together the
article.
Conflict of interest The authors declare no competing interests.
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