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Apex predators are crucial for maintaining ecological patterns and processes, yet humans hinder their ability to fulfil this role by displacing them from the landscape. Many apex predator species such as African lions (Panthera leo) are experiencing catastrophic declines as a result of competition with growing human populations. Increasing our understanding of the competitive interactions between lions and humans, as well as identifying thresholds of lion tolerance to human activities are important both for lion conservation and our understanding of apex predator ecology in the Anthropocene. We investigated the relative and cumulative influences of anthropogenic pressures on lion occurrence across a 73 000 km2 multi-use landscape in southern Africa. We developed occupancy models from replicated detection/non-detection spoor surveys across gradients of anthropogenic and biotic features. We tested the two hypotheses that African lions were most limited by 1) interference competition with humans or 2) exploitative competition with humans and evaluated the relative contribution of individual anthropogenic and biotic variables to lion occurrence. Our models predicted that lions occupied 49% of the landscape. The strongest determinants of lion occupancy were negative associations with pastoralism and bushmeat poaching, and a positive association with preferred prey. Thus, lions in this landscape are limited by a combination of interference and exploitative competition with poachers and pastoralists. However, interference competition with pastoralism was the biggest driver limiting lion occupancy, with a clear disturbance threshold for lions cumulating in a near complete loss of lions from the landscape when cattle surpass 21% occurrence. This study provides a predictive understanding of the top-down impacts of humans on the world's vulnerable apex carnivores.
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Original Research Article
Africa's apex predator, the lion, is limited by interference and
exploitative competition with humans
Kristoffer T. Everatt
, Jennifer F. Moore
, Graham I.H. Kerley
Centre for African Conservation Ecology, Department of Zoology, Nelson Mandela University, South Africa
Panthera, USA
University of Florida Department of Wildlife Ecology and Conservation, USA
article info
Article history:
Received 27 March 2019
Received in revised form 15 August 2019
Accepted 16 August 2019
Apex predators are crucial for maintaining ecological patterns and processes, yet humans
hinder their ability to full this role by displacing them from the landscape. Many apex
predator species such as African lions (Panthera leo) are experiencing catastrophic declines
as a result of competition with growing human populations. Increasing our understanding
of the competitive interactions between lions and humans, as well as identifying thresh-
olds of lion tolerance to human activities are important both for lion conservation and our
understanding of apex predator ecology in the Anthropocene. We investigated the relative
and cumulative inuences of anthropogenic pressures on lion occurrence across a
73 000 k m
multi-use landscape in southern Africa. We developed occupancy models from
replicated detection/non-detection spoor surveys across gradients of anthropogenic and
biotic features. We tested the two hypotheses that African lions were most limited by 1)
interference competition with humans or 2) exploitative competition with humans and
evaluated the relative contribution of individual anthropogenic and biotic variables to lion
occurrence. Our models predicted that lions occupied 49% of the landscape. The strongest
determinants of lion occupancy were negative associations with pastoralism and bush-
meat poaching, and a positive association with preferred prey. Thus, lions in this landscape
are limited by a combination of interference and exploitative competition with poachers
and pastoralists. However, interference competition with pastoralism was the biggest
driver limiting lion occupancy, with a clear disturbance threshold for lions cumulating in a
near complete loss of lions from the landscape when cattle surpass 21% occurrence. This
study provides a predictive understanding of the top-down impacts of humans on the
world's vulnerable apex carnivores.
©2019 The Authors. Published by Elsevier B.V. This is an open access article under the CC
BY-NC-ND license (
1. Introduction
Apex predators have important ecological roles, exerting disproportionate inuence over the structure and function of
ecosystems, such that the consequences of their disappearance can reverberate through trophic levels (Estes et al., 2011;
Ripple et al., 2014). Apex predators are extinction prone due to the rarity imposed on them by the energetic constraints of
their trophic position (Carbone and Gittleman, 2002;Cardillo et al., 2004). However, despite this natural rarity many apex
*Corresponding author. Centre for African Conservation Ecology, Department of Zoology, Nelson Mandela University, South Africa.
E-mail address: (K.T. Everatt).
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Global Ecology and Conservation 20 (2019) e00758
predators are in competition with humans (Treves and Karanth, 2003;Ripple et al., 2014). In the age of the Anthropocene,
these competitions have now pushed most species of apex predators into a conservation crisis (Dirzo et al., 2014;Ripple et al.,
2014). Thus,there is a need for a betterunderstanding of the nature of these competitiveinteractions including distinguishing
the relative effects of interference and exploitative competition on apex predator conservation.
For instance, African lions (Panthera leo) exert inuence on lower trophic levels (Estes et al., 2011;Tambling et al., 2012),
with complex cascading effects (Le Roux et al., 2018) and are in competition with humans (Kissui, 2008;Khorozyan et al.,
2015). Lions across Africa have suffered an estimated 50% decline in abundance and 75% reduction in range over the past
20e50 years (IUCN, 2006:Riggio et al., 2012;Bauer et al., 2015). Many populations continue to decline (Bauer et al., 2015). The
proximate cause of declining African lion populations is increasing competition with humans. Interference competition can
take the forms of persecution of lions in retaliation to depredation or perceived depredation of livestock (Woodroffe and
Frank, 2005), unsustainable trophy hunting of lions (Loveridge et al., 2007), poaching (illegal hunting) of lions (Everatt
et al., unpublished) or behavioural exclusions (Oriol-Cotterill et al., 2015;Smith et al., 2017). Exploitative competition, where
resources required by lions are limited by humans, include habitat loss to expanding agricultural, settlement or other human
land uses (Riggio et al., 2012) and the loss of wild prey resources to hunting by humans for meat (Lindsey et al., 2013).
Large carnivore species differ in their degree of competitive interactions with humans (Cardillo et al., 2004) and therefore
in their ability to persist in human-impacted landscapes. For instance, leopards (Panthera pardus) and cougars (Puma con-
color), both mid-sized, elusive and solitary generalists, can exist at relatively high densities even in unprotected human-
dominated landscapes by subsisting on domestic animals or smaller prey (Athreya et al., 2013;Moss et al., 2016). In
contrast, the abundance of grizzly bears (Ursus arctos), a larger carnivore which is more prone to conicts with humans is
more limited by human disturbance (Apps et al., 2004;Linke et al., 2013). Lions are among the largest of terrestrial carnivores,
are more gregarious and less cryptic then other felids, and display a preference for larger prey species whose weight category
overlaps with domestic livestock and wild species also sought after by human hunters (Schaller, 1972;Hayward et al., 2007).
Lions are also dominant over sympatric apex predators, including leopards, cheetah (Acinonyx jubatus), spotted hyenas
(Crocuta crocuta) and African wild dogs (Lycaon pictus)(Palomares and Caro, 1999;Durant, 2000) and hence unlikely to have
evolved appropriate strategies for mitigating competition with syntopic dominant predators. Lions are also larger and more
vocal, making their presence more obvious than the other syntopic predators. These characteristics may increase their
competitive interactions with humans relative to other apex predators and make humans less inclined to tolerate their
presence, thus reducing their ability to persist in anthropogenically-impacted landscapes.
In this study, we explore the roles of interference and exploitative competitive interactions between lions and humans,
and specically the relative importance of the various anthropogenic pressures on lion occurrence across the landscape. We
tested the hypotheses that a) lions were most limited byinterference competitionwith humans or b) lions were most limited
by exploitative competition with humans. We predicted that if lions were more limited by interference competition with
humans, then the occupancy of lions would be best predicted by a negative relationship with pastoralism and poaching
activities, and if lions were more limited by exploitative competition with humans then the occupancy of lions would be best
predicted by a positive relationship with prey abundance and a negative relationship with bushmeat poaching. Additionally,
we sought to identify thresholds of occurrence for lions along an increasing gradient of limiting anthropogenic pressures.
Thus, we add to the body of knowledge about the effects of competition by (1) describing anthropogenic effects on apex
predator ecology, and (2) providing information on competitive abilities of lions, which is crucial for predicting habitat
suitability and determining conservation needs of lions.
2. Methods
2.1. Study area and population
This study was carried out between 2014 and 2016 in the Greater Limpopo Lion Conservation Unit (GLLCU) of South Africa,
Mozambique and Zimbabwe (IUCN, 2006). The GLLCU covers approximately 73 00 0 km
, and includes South Africa's Kruger
National Park (KNP) (19 485 km
), Zimbabwe's Gonarezhou National Park (GNP) (5 053km
), and Mozambique's Limpopo
National Park (LNP) (11 233 km
), Banhine National Park (BNP) (7 250 km
) and Zinave National Park (ZNP) (4 000 km
several private game reserves, community hunting, grazing, farming and logging areas (Fig. 1). The study area includes a
relatively discrete lion population whose distribution can be dened by a hard edge along the fenced western boundary of the
greater KNP and the fenced northern boundary of GNP, and the eventual effective absence of lions along the eastern and
southern reaches of the GLLCU (this study).
Based on the large amount of available lion habitat and the size of the lion population, the GLLCU is considered one of
Africa's ten remaining population strongholds (Riggio et al., 2012). At the onset of this study (2014), KNP supported a stable
and protected population of >1600 lions (Ferreira and Funston, 2010), and GNP, LNP and BNP supported small populations
estimated at 33, 66 and 10 lions, respectively, with the latter three all below their respective ecological carrying capacities
(Groom et al., 2014;Everatt et al., 2014; K. Everatt, unpublished). Throughout the majority of the Mozambican portion of the
GLLCU lions are faced with the compounded challenges of prey depletion caused by bushmeat poaching, mortalities as by-
catch in bushmeat snares, poaching for body parts, retaliatory or pre-emptive killing by pastoralists and loss of habitat to
agricultural land conversion (Everatt et al., 2014; K. Everatt, unpublished). All three of the Mozambican National Parks are
inhabited by people who practice a variety of subsistence activities (cropping, pastoralism, hunting and gathering) as well as
K.T. Everatt et al. / Global Ecology and Conservation 20 (2019) e007582
high levels of commercial poaching of ungulates for bushmeat and of elephants (Loxodonta africana), rhino species (Cera-
totherium simum and Diceros bicornis) and lions for body parts (Everatt et al., 2015;Everatt et al., 2016; Everatt et al., un-
published; this study). Vegetation across the region is classied as mixed savanna, woodland and grasslands (Stalmans et al.,
2004). Wild ungulate densities across the GLLCU are spatially heterogeneous, largely due to the heterogeneous nature of
wildlife protection. For instance, impala (Aepyceros melampus) occur at densities >8/km
in KNP, 1.21/km
in GNP, 0.10/km
LNP and 0.07/km
in BNP (Redfern et al., 2002;Dunham et al., 2010;Grossmann et al., 2014;Stalmans and Peel, 2009) and
buffalo (Syncerus caffer) at densities of 1.50/km
in KNP, 0.46/km
in GNP, 0.12/km
in LNP and 0.002/km
in BNP (Seydack
et al., 2012;Dunham et al., 2010;Grossmann et al., 2014;Stalmans and Peel, 2009).
Fig. 1. Location of the study area (dark area in inset) in southern Africa and the survey area overing the Greater Limpopo Lion Conservation Unit of South Africa,
Mozambique and Zimbabwe, with the overlaid grid of 200 km
cells. Grid cells that were randomly selected for sampling are shaded.
K.T. Everatt et al. / Global Ecology and Conservation 20 (2019) e00758 3
2.2. Study design
Interspecic competition can be inferred by examining the dynamics of multi-species' occupancy assumed to be resultant
of one speciesnumerical or behavioural response to the presence of another (Linnell and Strand, 2000;Haynes et al., 2014).
We examined the heterogeneity in lion occupancy in relation to gradients of wild prey, domestic livestock and bushmeat
poaching pressures across the Greater Limpopo Lion Conservation Unit of southern Africa. In this study we dene bushmeat
poaching as the sum of all illegal or unregulated hunting of wildlife for meat (see Appendix 1 for examples). There was no
legal regulated hunting occurring in our study area during the surveys. We developed single-season occupancy models from
spatially replicated presence-absence walking spoor surveys of lions, their prey and threats, with an interest in two pa-
rameters, the probability of occupancy (
) and the probability of detection (p). Hierarchical modelling of covariates was used
to make inferences on the ecological factors limiting lion occupancy and detectability (MacKenzie et al., 2002). We made the
following assumptions for the estimator occupancy ѱto be interpreted as the proportion of area occupied: 1) sites were
closed to changes in occupancy; 2) species were not falsely identied; 3) detections were independent, or, if not then such
dependencies were modelled (Hines et al., 2010); and 4) all heterogeneity in occupancy or detection probability was modelled
using covariates (MacKenzie et al., 2002). To estimate the proportion of area occupied by lions, our sample units (sites) were
dened as 200 km
grid cells. Lion prides are known to occupy home ranges of between 30 km
(Kissui et al., 2009) and 1
450 km
(Funston, 2011) in size, with the size inversely related to the availability of prey (Van Orsdol et al., 1985). The average
home range in southern Kruger National Park, an area in the region with some of the highest relative prey densities (Ferreira
and Funston, 2010), is 100 km
(Funston et al., 2003). We therefore assumed that using grid cells of twice this size would be
large enough to reduce spatial autocorrelation between sites but small enough to assume that entire grid cells are occupied,
thus reducing the chance of occupancy over-estimation. A grid of 14.142 km 14.142 km (200 km
) cells was placed over the
entire landscape and 150 of these were randomly chosen for sampling, excluding any cells that consisted of over 50% set-
tlement areas. The resulting survey area covered the full gradients of anthropogenic disturbances and wild prey and livestock
densities from across the GLLCU (Fig. 1). A second level of randomness was incorporated by dividing each grid cell into four
quadrats (50 km
) and selecting one of these for obligate sampling.
In order to reduce the likelihood of changes in occupancy during sampling (closure) individual grid cells were sampled
within 36-h periods and adjoining grid cells were surveyed in succession (Hines et al., 2010). We acknowledge that a single
survey at this landscape scale however does not consider seasonal variations. Surveys were conducted by the same expe-
rienced tracker. All animal detections were represented by unambiguously identied tracks and bushmeat poaching de-
tections were identied by unambiguously identied sign, including snares, traps, shotgun shells, butchered carcasses, active
poaching camps and encounters with poachers (see Appendix 1)(Stuart and Stuart, 2013). Within each 200 km
cell an
average of 40 sequential 1 km detection non-detection samples were walked searching for lion tracks, other animaltracks and
bushmeat poaching sign, with at least one sample reaching into the randomly selected quadrat and the rst sample selected
upon access to the cell (Karanth et al., 2011;Thorn et al., 2011;Whittington et al., 2015). While this approach had not yet been
used for lions the survey effort used here was comparable to the required survey effort determined by Karanth et al. (2011) for
sampling the landscape occupancy of tigers (Panthera tigris). Due to poor tracking substrate and a lack of roads throughout
much of the study area, all transects were surveyed on foot. Sampling transects were chosen to maximize detection of rare
carnivores by searching along game trails and habitat edges, thereby minimizing the likelihood of false absences (MacKenzie
et al., 2006).
As an apex predator, lions are naturally limited by bottom up resources (Hayward et al., 2007), but also by top-down
anthropogenic pressures (Everatt et al., 2014) due to their wide exposure to human-driven ecological processes (Dirzo
et al., 2014). In order to model heterogeneity in lion occupancy, we collected covariate data for each 1 km sample which
we assumed to be biologically relevant to lion habitat use at the home range scale (Mitchell and Hebblewhite, 2012). These
covariates included the presence/absence of wild prey species, domestic livestock (cattle) and bushmeat hunting, and the
landscape variables (Table 1,Appendix 2). We combined important lion prey species into two categories: Preferred prey,
with the species preferentially selected by lions across Africa and in our study area and All preywith the combination of all
wild ungulates species which are predated on by lions in our study area (including preferred prey species) (Hayward and
Kerley, 2005; K Everatt, unpublished) (See Appendix 1 for the full list of species). The landscape variables that we
assumed to be important in determining lion occupancy (the distance from the centre of Kruger and Gonarezhou National
Parks (areas offering greater wildlife protection), the distance from settlement edges and the amount of riparian habitat) were
extracted for each 1 km sample from raster layers ( in ArcGIS 10.3.1 ( 1).
The variables that may inuence the detectability of tracks in this landscape include the observer skill, recent rains, angle
of sun, substrate quality and livestock and human densities (Funston et al., 2010;Stuart and Stuart, 2013). We approached the
sampling such that these biases were minimized or modelled as covariates. Surveys were conducted at least four days post
heavy rains, and surveys were not conducted at mid-day when the angle of the sun makes detecting tracks more difcult. In
addition, we rated the quality of substrate foreach 1 km sample as good, medium or poor based on the likelihoodof being able
to see a carnivore track if it was there and recorded the transect type. These were then considered as detection covariates
along with relative livestock and human impacts (Table 1). All detection/non-detection data and transect positions were
recorded by the Cyber tracker ( program on a smart phone.
K.T. Everatt et al. / Global Ecology and Conservation 20 (2019) e007584
2.3. Analytical methods
2.3.1. Occupancy modelling
Site occupancy
and detection probability pwere estimated using the maximum likelihood functions (MacKenzie et al.,
2006) of the single season correlated detections option in PRESENCE v. 11.6 (Hines, 2006). The correlated detections model
deals with the positive spatial autocorrelation of detections when sampling is not random with replacement and the
detection in one sample translates to an increased probability of detection in the next sample. Such situations are faced when
sampling for a mobile species along a trail (Hines et al., 2010). General notations for the correlated detections model are
provided in Hines et al. (2010). Continuous site covariates were standardized using a z-scale. Multi-collinearity between
variables was tested for using a Pearson correlation coefcient with an exclusion criterion of r >0.6 in MS Excel v. 1906 (Clare
et al., 2015).
2.4. Model selection procedures
We were interested in the relative contribution of anthropogenic (bushmeat poaching, pastoralism, protected area
management) and natural covariates (preyavailability) on the landscape-scale occupancy of lions. Models were ranked based
on Akaike Information Criterion (AIC) (Hines et al., 2010) and considered tobe strongly supported if
AIC 2. A candidate set
was developed which considered all models with
AIC ˂7 whose combined model weights 0.95, excluding the models that
did not reach numerical convergence. AIC weights were used to determine the weight of evidence for each model and they
were summed for each covariate in the 95% condence set (Burnham and Anderson, 2002). Variables with high summed
model weights were considered more important in explaining heterogeneity in lion occupancy. The sign of the
was used to determine the direction of inuence of covariates (MacKenzie et al., 2006). Covariates were considered robust if
the 90% condence interval
±1.65 standard error SE did not include zero (Burnham and Anderson, 2002). Overall
parameter estimates for detection probability pand occupancy ѱwere calculated using a weighted model averaging tech-
nique (Burnham and Anderson, 2002).
2.5. Identication of ecological thresholds
We were interested in identifying ecological thresholds in lion occupancy in relation to important natural and anthro-
pogenic variables. We plotted the model averaged estimate of lion occupancy against the highest ranking signicant
explanatory variables. Using the segmentedpackage in R (R Core Team, 2017)wet piecewise broken stickregression
models to the data to identify breakpoints where the relationship between the response and explanatory variables changes to
estimate ecological thresholds (Toms and Villard, 2015).
Table 1
Variables assumed to describe lion occupancy and detectability, their ecological relevance and metrics recorded from each 1 km samples of spoor surveys or
from GIS maps.
Variable Ecological relevance Metric recorded Source
Species preferentially selected by lions across Africa and in our study area (Hayward and
Kerley, 2005; K. Everatt, unpublished)
Summed presence/absence of each
All prey All prey species taken by lions in our study area (Hayward and Kerley, 2005; K. Everatt,
Summed presence/absence of each
Direct persecution of lions and/or depletion of lion prey Presence/absence Spoor
Cattle Persecution of lions by pastoralists/potential lion prey Presence/absence Spoor
Increased protection of lions and prey Distance (km) from centre of Kruger
of Gonarezhou NP
Settlement Increased human impact Distance (km) from nearest
settlement edge
Habitat feature known to be selected by some lion prey (Estes, 1991) and selected by
lions (Hopcraft et al., 2005)
Number of 30 30 m riparian pixels
per grid cell
Substrate Inuence on the detectability of tracks. Scoring based on assumed likelihood of detecting
lion tracks if lions were present.
Good/medium/bad Spoor
Transect type Lions may show selection/avoidance of transect types Road/track/trail/river/bush Spoor
See Appendix 1 for full description of prey species and bushmeat poaching sign recorded.
K.T. Everatt et al. / Global Ecology and Conservation 20 (2019) e00758 5
3. Results
3.1. Occupancy by lions
A total of 3759 1 km sampling occasions were walked across 103 grid cells (range 10e45 km/cell, mean ¼34.5 km/cell)
resulting in 26 000 km
of sampled habitat. Lions were detected in 215, poaching in 485, cattle in 783 and one or more of
preferred prey species in 2608 of the samples (Appendix 3).
The covariate combination of Preferred preyand All prey(r ¼0.81) as well as All preyand Cattle(r ¼0.71) were
found to be correlated and were not included in models together leaving 24 candidate models of statistically uncorrelated
variables (Appendix 4).
3.2. Model selection
We rst accounted for the probability of detection pby ranking all covariates for pincluding the two detection only
covariates (see Table 1). The top ranking model included the covariates substrate, cattle occurrence, and distance to nearest
settlement on the probability of detection (
,th0(),t.h1(.),p(Sub. þCþS),th0pi(.);Table 2). Given our sampling effort, the
probability of detecting lions in a grid cell from the top-ranking model was calculated at b
p¼0.344 ±SE ¼0.092. This model
was then held constant for all further analysis and only the covariates describing occupancy ѱwere left to vary. We then
compared 24 models describing the inuence of the covariates, in isolation and in combinations, on lion occupancy. From all
top-ranking models, the weighted average occupancy estimate was b
¼0.488 ±SE ¼0.079.
Heterogeneity in the occupancy of lions was best explained by the signicantly negative relationship with the occurrence
of cattle, which occurred in all the highest-ranking models (
¼16.486, SE ¼8.788, w¼1.0). Lion occupancy was also
explained by the signicantly positive relationship with the occurrence of preferred prey species (
¼1.199, SE ¼0.723,
w¼0.516). A negative relationship with bushmeat poaching (
¼0.455, SE ¼0.725, w¼0.753) and positive relationship
with proximity to protected area centres (
¼0.021, SE ¼0.039, w¼0.178) also contributed to explaining lion occupancy
(Table 2,Fig. 2).
3.3. Identication of ecological thresholds
A negative non-linear response was identied between the weighted average estimates of lion occupancy b
and the
occurrence of cattle with a threshold at the value of cattle occurrence of 0.208 (Fig. 3a). Mean site estimates were b
¼0.677 ±SE ¼0.105 at sites with cattle occurrence values of less than 0.208 (74 sites) and were b
¼0.005 ±SE ¼0.013 at
sites with cattle occurrence values of greater than 0.208 (29 sites). A positive non-linear response was identied between lion
occupancy and the occurrence of preferred lion prey with a threshold occurring at the value of preferred prey occurrence of
0.218 (Fig. 3b). Mean site estimates were b
¼0.063 ±SE ¼0.012 at sites with preferred prey occurrence values of less than
0.218 (11 sites)and were b
¼0.539 ±SE ¼0.087 at sites with preferred prey occurrence values of greater than 0.218 (92 sites).
The relationship between lion occupancy and the occurrence of bushmeat poaching displayed a weak negative correlation of
¼0.0939 (Fig. 3c).
4. Discussion
Many apex carnivore species now nd themselves in a conservation crisis, being limited through competitive interactions
with humans (Ripple et al., 2014). In this study we sought to identify what limits lions in human-impacted landscapes by
considering the relative roles of interference and exploitative competition between lions and humans, and the relative
importance of different anthropogenic pressures on lion occupancy. We further identied important thresholds for the oc-
cupancy of lions across the landscape.
Table 2
The summary of the best models describing the effects of covariates on lion occupancy ѱand detectability p. Number of sites ¼109. Covariates include Cattle
(C), Preferred prey (PP), Bushmeat poaching (BP), Protected areas (PA), Settlements (S), Substrate (Sub) and Transect type (T).
AIC AIC weight Model likelihood Number of parameters
,th0(.),th1(.),p(Sub þCþS),th0pi(.) 0.00 0.952 1 8
,th0(.),th1(.),p(Sub þCþSþT),th0pi(.) 5.99 0.048 0.4795 7
(C þPP), th0(.),th1(.),p(Sub þCþS),th0pi(.) 0.00 0.3180 1 10
(C),th0(.),th1(.),p(Sub þCþS),th0pi(.) 0.74 0.2196 0.6907 9
(C þPP þBP),th0(.),th1(.),p(Sub þCþS),th0pi(.) 1.60 0.1429 0.4493 11
(C þBP),th0(.),th1(.),p(Sub þCþS),th0pi(.) 1.96 0.1193 0.3753 10
(C þPA),th0(.),th1(.),p(Sub þCþS),th0pi(.) 2.45 0.0934 0.2938 10
(C þBP þPP þPA),th0(.),th1(.),p(Sub þCþS),th0pi(.) 3.50 0.0553 0.1738 12
K.T. Everatt et al. / Global Ecology and Conservation 20 (2019) e007586
We found that lions occupied approximately 49% of the full 26 0 00 km
area of sampled habitat. The presence of cattle was
the single strongest explanatory variable of lion occupancy, with the cattle covariate appearing in all the top-ranking models
and having a signicant negative inuence on lion occurrence. Because lions are known to predate on cattle (Kissui, 2008;
Valeix et al., 2012;Tumenta et al., 2013;Loveridge et al., 2017) this signicant negative relationship between cattle presence
and lion occupancy likely indicates that the costs associated with cattle depredation, including persecution by pastoralists,
must be higher than the energetic rewards gained by lions byconsuming cattle. It is important to note that the occurrence of
cattle is used here as a proxy for the impacts of likely retaliatory actions taken towards lions by cattle owners.
The inuence of cattle on lion occupancy could indicate that lions are individually aware of the risks associated with
occupying pastoralist areas and exhibit a behavioural avoidance of these landscapes or of threats within these landscapes
(Oriol-Cotterill et al., 2015;Loveridge et al., 2016;Suraci et al., 2019), or that lions are excluded from cattle areas through
persecution of lions by pastoralists (Woodroffe and Frank, 2005;Kissui, 2008). Behaviour avoidance of high-risk areas is a
common strategy among species under predation pressure (Thaker et al., 2011) and even apex predators are increasingly
having to adjust their behaviour in response to top-down pressures exerted by humans (Oriol-Cotterill et al., 2015;Smith
et al., 2015,2017). Alternatively, the inuence of cattle on lion occupancy documented here could also indicate a numeri-
cal response where lions are excluded from cattle-occupied lands through higher levels of persecution. An analysis of the
sources of lion mortalities in our study area found that 41.2% of documented lion mortalities were attributed to persecution by
pastoralists (K Everatt, unpublished), conrming that at least some of the heterogeneity in lion occupancy described here
should be attributed to a numerical response to this anthropogenic pressure rather than a behavioural response.
Large carnivores are known to respond to thresholds of human density beyond which the species or populations become
locally extinct (Woodroffe, 2000). We found a non-linear continuous response between lion occupancy and cattle occurrence
cumulating in a near absence of lions at sites with cattle occurrence greater than 21%. This threshold indicates a clear point
beyond which the relative occurrence of cattle and associated retaliatory or pre-emptive killing of lions for livestock
depredation prohibits the occurrence of lions in the landscape. In this case the 21% cattle occurrence threshold can be
interpreted as meaning that 21% of a given area is currently being used by cattle (within an approximate 14-day window (see
methods)). We interpret the resultant low occupancy rate of lions in cells with more than 21% cattle occurrence to mean that
lions are only able to occasionally use a landscape with this level of pastoralist impact. Beyond this level of pastoralist impact
lions may disperse into such areas but are unlikely to become resident; an interpretation which is validated by the author's
observations that many of the lions found in livestock areas, and which are involved in livestock conict, were of dispersal age
and were often subsequently killed in retaliation of real or perceived livestock depredation (K Everatt, unpublished). Similar
human disturbance thresholds have been identied in other large predator systems including thresholds of resource
extraction beyond which grizzly bears cease to occur in western Canada (Lindsey et al., 2013) and thresholds of housing
density beyond which cougars cease to occur in the western US (Maletzke et al., 2017).
Knowledge of these thresholds is helpful for guiding conservation (Foley et al., 2015). For instance, assuming that the
response of lion occupancy to this threshold of cattle occurrence indicates a numerical response to persecution by pastoralists
Fig. 2. a) The relative contribution of important covariates (Pw>0.95), and b) the strengths and directions of the covariates explaining lion occupancy. The
covariates marked by * have
-coefcients with greater than 90% signicance.
K.T. Everatt et al. / Global Ecology and Conservation 20 (2019) e00758 7
and assuming that lions select for cattle as prey (Hayward and Kerley, 2005), we can predict that sites within this study area
that have greater than 20% cattle occurrence could be acting as ecological traps for lions in the region (Battin, 2004).
The disturbance threshold we report here builds on the substantial body of knowledge describing the inuences of
pastoralism on lion ecology (Valeix et al., 2012;Schuette et al., 2013;Oriol-Cotterill et al., 2015;Loveridge et al., 2016;Suraci
et al., 2019) and aligns with continent-scale meta-analyses indicating a negative inuence of pastoralism on lion conservation
(Packer et al., 2013;Lindsey et al., 2017).
These data provide evidence of the negative impact of pastoralist activities on lion occupancy considering the response of
lions across a spatial gradient of anthropogenic pressures, analogous to a response of increasing pressures at a single site. This
space for timeapproach is useful when studying ecological relationships between long-lived species which exhibit slower
and thus more difcult to detect responses to environmental changes (Estes et al., 2010). The response of lions to changes in
cattle occurrence identied here could help predict the species' response to Africa's increasing cattle herds and pastoralist
footprint (Wittemyer et al., 2008). It is interesting to note, however, that because our data originates from surveys across the
spatial gradients in a time snapshot it is possible that the occupancy threshold of lions in relation to cattle may be hysteretic,
that is being dependent on the direction of change in cattle occurrence (Scheffer et al., 2001). Cattle in our study area are
mostly restricted to Mozambique, where the abundance of cattle has generally been on the increase following economic
recovery after the end of country's civil war (Grossmann et al., 2014) and it is therefore possible that the threshold of lion
occurrence might be placed at much lower cattle occurrence in a system that is moving from higher to lower cattle impacts
(see Estes et al., 2010). For instance, a cattle-disturbance threshold in parts of East and West Africa may be expected to be
lower due to extreme degradation by overgrazing coupled with climate change causing, at least localized, declines of humans
and livestock (Wittig et al., 2007;Mganga et al., 2018). A successful carnivore-livestock conict mitigation scheme (for
example Hazzah et al., 2014;Lichtenfeld et al., 2015) that reduces the number of lions killed by pastoralists in retaliation to
depredation could also shift the disturbance thresholds shown here.
The occupancy of lions across our study area had a signicant positive relationship with the occurrence of their preferred
prey. The importance of the availability of prey resources for determining apex carnivore habitat use is well known and it is
expected to be the most important variable in the natural functioning of predator-prey systems (Hopcraft et al., 2005;
Mitchell and Hebblewhite, 2012;Everatt et al., 2015). Interestingly, the relationship between lion occupancy and the
occurrence of their preferred prey was nonlinear with a stronger relationship found at sites below a threshold of
Fig. 3. Relationships between the weighted average estimates of lion occupancy and the top-ranking signicant explanatory covariates; a) cattle, b) preferred
prey and c) bushmeat poaching occurrence.
K.T. Everatt et al. / Global Ecology and Conservation 20 (2019) e007588
approximately 20% preferred prey occurrence. This stronger response may indicate the absence of density-dependant social
mechanisms regulating lion occupancy in these sites and indicates lion population or behavioural instability at these sites.
While the occupancy of lions was positively determined by their prey, it was also equally negatively determined by the
occurrence of bushmeat poaching activities (each contributing to 75.3% of the combined model weights). Prey populations
are severely depleted throughout the Mozambique portions of the study area where bushmeat poaching is widespread
(Everatt et al., 2019;Lindsey et al., 2017;Baghai et al., 2018). For instance, wild ungulate biomass was estimated at less than
20% of carrying capacity in the Limpopo National Park (Baghai et al., 2018) where bushmeat poaching occurred across 80% of
sites surveyed (Everatt et al., 2014). Everatt et al. (2014) showed that bushmeat poachers in Limpopo National Park select for
areas closer to settlements (as they typically travel by foot) and with medium-sized ungulates. Areas closer to settlements in
this landscape have become denuded of most wild ungulates other than the smallest of species such as common duikers
(Sylvicapra grimmia) (K Everatt, unpublished) which fall outside of lion's preferred prey range. The inuence of the bushmeat
poaching variable on the occupancy by lions in this study area likely includes lions' selection against these prey-depleted
lands. The impact of competition with bushmeat poachers for prey on the occurrence of an apex predator would be ex-
pected to differ according to the degree of niche overlap between the predator and humans. For instance, a difference in
competitive ability with humans has been documented between tigers and leopards in India (Athreya et al., 2013). In Africa,
leopards which exploit smaller species (Hayward et al., 2007) could be expected to be more successful in wildlife depleted
landscapes than lions. A previous analysis found lion habitat use in Limpopo National Park to be strongly negatively asso-
ciated with bushmeat poaching activities (Everatt et al., 2014), while in contrast, leopard habitat use in the same area and time
was positively associated with both bushmeat poachers and lions (Strampelli et al., 2018). It is likely, however, that leopards
were not selecting for areas with lions or bushmeat poachers per se but rather selecting for the same hunting opportunities as
lions and bushmeat poachers. However, the bushmeat poaching variable in this study may also indicate other drivers in
addition to the depletion of prey. In this system, bushmeat poachers are often accompanied by large packs of domestic dogs
whose presence can be avoided by lions. Domestic dogs have been used successfully to deter lions from livestock (Bauer et al.,
2010). Dogs are loud, mobile and conspicuous, all of which are characteristics of a threat that would be relatively easy to
detect and avoid. Avoiding bushmeat hunters who use snares and traps, which are also commonly employed hunting tools in
this study area, however, would be very different in that respect from avoiding hunters who use dogs. Snares and traps are set
and left in such a way that they are meant to remain undetected by wildlife and it would thus be more difcult for lions to
adapt an avoidance behaviour to this type of cryptic activity. In fact, during this and other surveys in the Limpopo and Banhine
National Parks we have recorded lions scavenging fromsnares (K Everatt, unpublished) suggesting that lions do not recognize
this threat or show avoidance of snares.
The third explanation of a strong negative effect of bushmeat hunting on lion occupancy is that lions are persecuted and
spatially excluded by poachers. For instance, bushmeat poaching bycatch accounts for 18% of all known human-caused lion
mortalities in Limpopo National Park (K Everatt, unpublished). On some of the scavenging events mentioned above the lions
have themselves then been caught and killed in adjoining snares, suggesting that bushmeat poaching can also be acting as an
ecological trap to lions. There is also an overlap between bushmeat poaching and a rise in the targeted poaching of lions
where bushmeat poachers as well as rhino and elephant poachers are also increasingly poaching lions for body parts (K
Everatt, unpublished).
Apex carnivores are facing massive declines around the world as a consequence of their interactions with humans (Ripple
et al., 2014). The theory of intra-guild competition is applicable to understanding the conservation crisis facing apex carni-
vores, such as African lions, when humans are recognized as an apex predator. This study adds to the emerging body of
empirical results and a predictive understanding of the top-down impacts of humans on the world's vulnerable apex
We thank the following institutions for providing necessary permits: Administraç~
ao Nacional das
Areas de Conservaç~
South African National Parks, Zimbabwe Parks, Parque Nacional de Limpopo, Parque Nacional de Banhine, Parque Nacional de
Zinave, Kruger National Park, Gonarezhou National Park, Karangani Reserve and Maunge Conservancy. We thank Panthera,
the Natural Sciences and Engineering Research Council of Canada, Nelson Mandela University and Wilderness Foundation for
funding. We thank Leah Andresen and Eden Everatt for help in the eld and three anonymous reviewers for their input on the
Appendix A. Supplementary data
Supplementary data to this article can be found online at
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... Lions are thought to be especially vulnerable to anthropogenic pressures because they are less cryptic than other large carnivores (e.g. among the largest in size, social and prefer wild prey whose size overlaps with livestock; Everatt et al., 2019). They are typically the dominant predator within their ecosystem. ...
... Our goal was to understand how different landscape features impact lion space use and to inform practitioners of relevant conservation and human-lion conflict mitigation strategies that can be derived from our results. Previous studies found that lions typically avoid areas of high human presence and anthropogenic risk-including towns, highways and agricultural and livestock areas-over large spatial scales (Elliot et al., 2014;Everatt et al., 2019;Loveridge et al., 2017;. However, lions have also exhibited fine-scale temporal partitioning when avoidance of human activity is not possible . ...
... If lions select for prey irrespective of livestock presence (i.e. irrespective of a landscape of fear), then conflict mitigation strategies could focus on enhancing prey availability (Bauer et al., 2010;Everatt et al., 2019Everatt et al., , 2023. If livestock are being selected where there is equally accessible prey, then lion accessibility to livestock will need to be reduced (e.g. ...
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Conserving large carnivores requires protecting landscape spaces that encompass all spatiotemporal scales of their movement. Large carnivores normally roam widely, but habitat loss and fragmentation can constrain their movement in ways that restrict access to resources and increase encounters with humans and potential conflict. Facilitating carnivore population coexistence with humans across landscapes requires conservation plans informed by patterns of carnivore space use, particularly at the human–wildlife interface. We sought to understand lion space use in Laikipia, Kenya. We conducted a path‐selection function analysis using GPS collar data from 16 lions to assess patterns of space use across a range of spatial scales (sedentary to home range expanses; 0, 12.5, 25 and 50 km) and temporal scales (day, dusk, night and dawn). Path‐selection results were then incorporated into space use maps. We found that most landscape features influenced path‐selection at the broadest spatial scale (50 km), representative of home range‐wide movement, thereby demonstrating a landscape‐wide human impact on lion space use. We also detected sub‐diurnal variation in lion path‐selection which revealed limited space use during daylight hours and increased space use overnight. Our results highlight that optimal support for human–lion coexistence should be temporally adaptive at sub‐diurnal scales. Furthermore, spatial approaches to lion conservation may be better generalized at broad spatial scales so that land management plans can account for home range patterns in lion space use.
... We conducted vehicle-based spoor (track) transect surveys along roads, to collect detection/ non-detection presence/absence data on large carnivores across the landscape. Large carnivores perform extensive movements along road networks [54], and spoor-based surveys have been shown to be an efficient and effective method to collect data on large carnivores over vast landscapes, including for all of our study species [55][56][57]. ...
... We employed a spatially replicated occupancy sampling approach [47,[55][56][57]. Each transect was divided into 500 m segments, and we recorded whether sign of each species was detected (1) or non-detected (0) within each segment. ...
... Our study shows that the method holds significant promise to improve understanding of population-level mechanisms in African conservation landscapes, and we encourage similar studies elsewhere, particularly given the existing availability of sign-based detection/non-detection datasets [55,56,59,87,88]. We especially encourage further investigations into how the effects identified vary across gradients of anthropogenic impacts, and protected area management strategies. ...
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Interspecific interactions can be a key driver of habitat use, and must be accounted for in conservation planning. However, spatial partitioning between African carnivores, and how this varies with scale, remains poorly understood. Furthermore, most studies have taken place within small or highly protected areas, rather than in the heterogeneous, mixed-use landscapes characteristic of much of modern Africa. Here, we provide one of the first empirical investigations into population-level competitive interactions among an African large carnivore guild. We collected detection/non-detection data for an eastern African large carnivore guild in Tanzania’s Ruaha-Rungwa conservation landscape, over an area of ~45,000 km ² . We then applied conditional co-occupancy models to investigate co-occurrence between lion, leopard, and African wild dog, at two biologically meaningful scales. Co-occurrence patterns of cheetah and spotted hyaena could not be modelled. After accounting for habitat and detection effects, we found some evidence of wild dog avoidance of lion at the home range scale, and strong evidence of fine-scale avoidance. We found no evidence of interspecific exclusion of leopard by lion; rather, positive associations were observed at both scales, suggesting shared habitat preferences. We found little evidence of leopard habitat use being affected by wild dog. Our findings also reveal some interspecific effects on species detection, at both scales. In most cases, habitat use was driven more strongly by other habitat effects, such as biotic resources or anthropogenic pressures, than by interspecific pressures, even where evidence of the latter was present. Overall, our results help shed light on interspecific effects within an assemblage that has rarely been examined at this scale. We also demonstrate the effectiveness of sign-based co-occurrence modelling to describe interspecific spatial patterns of sympatric large carnivores across large scales. We conclude by discussing the implications of our findings for large carnivore conservation in modern African systems.
... These behaviours suggest that lions may recognise the indirect risks associated with predating on a species which offers limited direct risks and that such responses to a pastoralistinduced landscape of fear may be a part of the optimal foraging of lions. Promoting lion persistence in prey-depleted and pastoralistimpacted landscapes may therefore ultimately be dependent on understanding the role of fear on lion selection or avoidance of livestock as prey (Khorozyan et al., 2015) and using this information to identify and maintain minimum thresholds of wild prey and space available to lions (Bauer et al., 2010;Everatt, Moore, et al., 2019;Khorozyan et al., 2015). ...
... Vegetation across the area is generally classified as mixed savannah woodland and grasslands (Stalmans et al., 2004). Wildlife populations are at or near carrying capacity in Kruger National Park and generally decline eastwardly due to management history and environmental variables (Everatt, Moore, et al., 2019). The majority of the Mozambican section of the study area is occupied by free-ranging cattle (Everatt, Moore, et al., 2019). ...
... Wildlife populations are at or near carrying capacity in Kruger National Park and generally decline eastwardly due to management history and environmental variables (Everatt, Moore, et al., 2019). The majority of the Mozambican section of the study area is occupied by free-ranging cattle (Everatt, Moore, et al., 2019). ...
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Optimal foraging and landscape of fear theories provide frameworks which can be useful for investigating animal's space and prey use decisions. Predators, such as African lions Panthera leo, are likely to respond to prey abundance, accessibility, profitability and potential risks, often anthropogenic in nature, while making foraging decision. Identifying the relative role of these processes has important conservation implications. We investigated the relative role of responses to a pastoralist landscape of fear within lion feeding and spatial ecology in a landscape at the human-wildlands interface. We collected spatial and predation data from 12 GPS-collared lions and ungulate count data from transects, along the South Africa-Mozambique border, including parts of Kruger, Limpopo and Banhine National Parks. We calculated Jacobs' Index values from 80 kills to investigate lion selection of wild and domestic ungulates as prey, used maximum entropy modelling to predict multi-season ungulate spatial occurrence and used resource selection functions to estimate the relative probability of use of wild and domestic ungulate areas by lions. All lions had access to wild prey and domestic livestock within their home ranges. Lions showed a strong selection for large-bodied wild ungulates as prey taking waterbuck, zebra, kudu and buffalo more frequently then predicted by their availability. Lions showed a slight avoidance of cattle as prey, with cattle outnumbering larger ungulates across much of the study area. Lions showed the greatest selection for habitats with high occurrences of wild prey, specifically areas with kudu, then nyala and buffalo, during the dry seasons and showed strong avoidance of cattle areas during the wet season; a season when cattle are kept closer to settlements and thus better protected and easier to predict and avoid. These results suggest that lions select for wild prey and habitats optimally, yet show a fear response to cattle and cattle areas. This duality in the foraging behaviour of lions suggests that efforts to mitigate human-lion conflict and preserve vulnerable lion populations should focus on both increasing wild ungulate populations as well as exploiting lion's fear of humans with careful consideration of the risks of livestock presence acting as an ecological trap for vulnerable lion populations. K E Y W O R D S African lion, carnivore conservation, habitat use, human-wildlife conflict, landscape of fear, optimal foraging, prey selection, resource-selection function 13652028, 0, Downloaded from
... Despite these trends, it remains poorly understood how anthropogenic pressures compare with biotic resources and interspecific effects in shaping space use and how these relationships vary among species (Balme et al., 2014;Everatt et al., 2019). Understanding the habitat requirements and tolerance limits of large carnivore communities in modern, multidimensional systems is critical for their conservation in increasingly heterogeneous and human-affected habitats (Balme et al., 2014). ...
... We employed a spatially replicated occupancy sampling approach (Everatt et al., 2019;Henschel et al., 2016;Petracca et al., 2019). Each transect was divided into 500-m segments, and we recorded whether a sign of each species was detected (1) or not detected (0) within each segment. ...
... For each large carnivore species, the prey species included as covariates were based on prey preferences described in the literature. The probability of illegal human activity was similarly modeled from our survey data and included as an explanatory covariate in the large carnivore models (Everatt et al., 2015(Everatt et al., , 2019. As for prey, modeling human activity explicitly from survey data rather than through proxies (e.g., distance to boundary and distance to ranger post) allowed us to quantify illegal human activity empirically, ensuring that the covariate was truly representative of anthropogenic disturbances within the PA complex. ...
Large carnivores increasingly inhabit human-impacted landscapes, which exhibit heterogeneity in biotic resources, anthropogenic pressures, and management strategies. Understanding large carnivore habitat use in these modern systems is critical for their conservation, as is the evaluation of competing management approaches and the impacts of significant land use changes. We employed occupancy modelling to investigate habitat use of an intact eastern African large carnivore guild across the 45,000 km2 Ruaha-Rungwa landscape, in south-central Tanzania. We determined the relative impact of biotic, anthropogenic, and management factors on five large carnivore species, at two biologically meaningful scales. We also specifically tested the effect of a novel trend of trophy hunting area abandonment on large carnivore occurrence. Our results reveal contrasting habitat use patterns: lion were found to be particularly vulnerable to illegal human activity, while African wild dog were instead limited by biotic features, avoiding areas of high sympatric predator density and using less-productive habitats. Spotted hyaena and leopard were able to persist in more disturbed areas, and across habitat types. There was no evidence of large carnivore occurrence being impacted by whether an area was used for photographic or trophy hunting tourism, with regular law enforcement being instead more important. All species fared better in actively managed hunting areas compared to those that had been abandoned by operators. Overall, our findings highlight the divergent habitat requirements within large carnivore guilds, and the importance of adopting an integrated approach to large carnivore conservation planning in modern systems. We also identified a novel threat to African conservation areas, in the form of decreased management investments associated with the abandonment of trophy hunting areas, and provide the first assessment of this significant land management change on a large carnivore population. Article impact statement: Habitat degradation associated with ongoing hunting area abandonment is shown to be a novel threat to large African carnivore populations. This article is protected by copyright. All rights reserved.
... Creation of effective protected areas for large carnivore conservation requires an understanding of seasonal factors that affect their habitat use 21,89 . Our findings on seasonal lion habitat use in relation to human and livestock density, distance to roads and rivers, and land cover were broadly consistent with previous research that demonstrated that lions balance seasonal patterns of prey availability 22,33 with avoiding humans 20,93 . We also quantified the importance of buffer protected areas and their strength of protection for lions. ...
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Protected areas that restrict human activities can enhance wildlife habitat quality. Efficacy of protected areas can be improved with increased protection from illegal activities and presence of buffer protected areas that surround a core protected area. Habitat value of protected areas also can be affected by seasonal variation in anthropogenic pressures. We examined seasonal space use by African lions (Panthera leo) within a core protected area, Serengeti National Park, Tanzania, and surrounding buffer protected areas with varying protection strengths. We used lion locations in logistic regression models during wet and dry seasons to estimate probability of use in relation to protection strength, distance to protected area edge, human and livestock density, distance to roads and rivers, and land cover. Lions used strongly protected buffer areas over the core protected area and unprotected areas, and moved away from protected area boundaries toward the core protected area when buffer protected areas had less protection. Lions avoided high livestock density in the wet season and high human density in the dry season. Increased strength of protection can decrease edge effects on buffer areas and help maintain habitat quality of core protected areas for lions and other wildlife species.
... During the last decade, the Markov model experienced an increasing use for sign-based surveys of large mammals moving along trails (e.g. Everatt et al., 2019;Karanth et al., 2011). Yet, its application remains scarce and still limited to few taxa and ecosystems. ...
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Assessing protected areas (PA) effectiveness for aquatic species is essential, as they are frequently recognised ineffective for freshwater ecosystems. By using spatially correlated replicates, the occupancy model with Markovian spatial dependence is well-suited to network-constrained environments. We applied this model to a semi-aquatic mammal, Galemys pyrenaicus, across the river network of the French Pyrenees. We found that occupancy is mainly influenced by climatic and hydrographic factors. Rainfall, forest cover and flow variability influence the overall high faeces detection. We then assessed the efficiency of the PA network to protect suitable streams for G. pyrenaicus by combining conservation gap analyses with two types of model outputs (i.e. occupancy probabilities and binary predictions). Using complementary indices and permutation tests, we found that about 25% of stream sections protected by PA are highly suitable for G. pyrenaicus and less than 5.5% of the most suitable sections benefit from a moderate to strong level of protection. Some highly suitable unprotected areas for G. pyrenaicus were identified where conservation measures should urgently be implemented. This study presents an innovative and integrative approach that opens future perspectives for development and additional applications to other taxa, difficult-to-survey or network-constrained environments.
... However, leopards feed on ungulate species targeted by wire-snare poachers (Henschel et al., 2011;Creel et al., 2018;Strampelli et al., 2018), and may scavenge on animals killed in snares (Strampelli et al., 2018). Snares are set in lines or clusters (Noss, 1998;Becker et al., 2013a), and predators that investigate struggling prey or carrion are susceptible to themselves becoming snared (Knopff et al., 2010;Everatt et al., 2019b). Therefore, despite the rarity of reports, leopards are presumably vulnerable to snaring like other large African carnivores. ...
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The impact of snaring and human-wildlife conflict (HWC) on large carnivore populations is of growing concern, and yet few empirical data are available. Mortality is the metric most often used, but non-lethal injuries that impact fitness are also important threats. However, because non-lethal injuries to wild carnivores are difficult to detect, they have received little study. Using straightforward forensic examination of the skulls of trophy-hunted lions and leopards from Luangwa Valley (LV) and Greater Kafue Ecosystem (GKE), Zambia, we identified non-lethal injuries consisting of snare damage to teeth and shotgun pellets in skulls. Wire snare entanglement can cause permanent, diagnostic damage to carnivore teeth when individuals bite and pull on the wire. Shotguns are used by poachers, as well as during HWCs to drive off carnivores perceived as threats. Carnivores struck by shotgun pellets can suffer non-lethal, but potentially toxic injuries such as pellets embedded in their skulls. Because poaching and HWC are generally more prevalent near human settlements, we predicted a higher incidence of anthropogenic injuries to carnivores in Luangwa where the human population is larger and more concentrated along protected area edges than Kafue. Contrary to expectation, anthropogenic injuries were more prevalent among lions and leopards in Kafue than Luangwa. Notably, definitive evidence of snare entanglement greatly surpassed previous estimates for these regions. Overall, 37% (41 in 112) of adult male lions (29% in Luangwa, 45% in Kafue) and 22% (10 in 45) of adult male leopards (17% in Luangwa, 26% in Kafue) examined had survived being snared at some point in their lifetime. Among adult male lions, 27% (30 in 112) had old shotgun pellet injuries to their skulls. Our procedure of forensic examination of carnivore skulls and teeth, some of which can be applied to live-captured animals, allows for improved detection of cryptic, non-lethal anthropogenic injuries. Further, our methods represent a consistent and economical way to track changes in the frequency of such injuries over time and between regions, thereby providing a direct measure of the effectiveness of conservation programs that seek to reduce poaching and HWC.
... This is especially the case if such efforts are accompanied by complementary landscape-scale indicators of status (e.g. sign-based occupancy surveys; Everatt et al., 2019;Henschel et al., 2020). ...
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1. Accurate and precise estimates of population status are required to inform and evaluate conservation management and policy interventions. Although the lion (Panthera leo) is a charismatic species receiving increased conservation attention, robust status estimates are lacking for most populations. While for many large carnivores population density is often estimated through spatially explicit capture-recapture (SECR) applied to camera trap data, the lack of pelage patterns in lions has limited the application of this technique to the species. 2. Here, we present one of the first applications of this methodology to lion, in Tanzania's Ruaha-Rungwa landscape, a stronghold for the species for which no empirical estimates of status are available. We deployed four camera trap grids across habitat and land management types, and we identified individual lions through whisker spots, scars and marks, and multiple additional features. 3. Double-blind identification revealed low inter-observer variation in photo identification (92% agreement), due to the use of xenon-flash cameras and consistent framing and angles of photographs. 4. Lion occurred at highest densities in a prey-rich area of Ruaha National Park (6.12 ± SE 0.94 per 100 km^2), and at relatively high densities (4.06 ± SE 1.03 per 100 km^2) in a community-managed area of similar riparian-grassland habitat. Miombo woodland in both photographic and trophy hunting areas sustained intermediate lion densities (1.75 ± SE 0.62 and 2.25 ± SE 0.52 per 100 km^2 , respectively). These are the first spatially explicit density estimates for lion in Tanzania, including the first for a trophy hunting and a community-managed area, and also provide some of the first insights into lion status in understudied miombo habitats. 5. We discuss in detail the methodology employed, the potential for scaling-up over larger areas, and its limitations. We suggest that the method can be an important tool for lion monitoring and explore the implications of our findings for lion management. 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.
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Human-wildlife conflict is a challenging issue that requires the attention of conservationists worldwide. Habitat fragmentation and encroachment reduce the abundance of prey species, and an increase in the number of predators leads to a higher risk of conflict with large cats such as leopards, jeopardizing conservation efforts. This study explored the spatio-temporal pattern of the human-leopard conflict in Bardia National Park, Nepal, from 2000 to 2020. To analyze the conflict with leopards, we used data (compensation cases filed in the park) from the buffer zone management office, the National Trust for Nature Conservation (NTNC), and the Department of National Park and Wildlife Conservation (DNPWC). Leopard attacks on livestock are increasing exponentially, with 3335 livestock killed in 2652 attacks occurring during the study period. Although livestock depredation by leopards occurred all over the park, the southern cluster has most documented livestock damage (64.01%). The eastern and northern clusters reported fluctuating and dispersed predation events, respectively. Our spatial analysis indicated no effect of topography (slope) on livestock depredation by leopards. We recorded the highest number of leopard attacks and predation during the dry winter season when the nights are longer and livestock remain in their sheds. This carnivore mostly limited its prey to small-sized livestock (95.77%) such as goats, sheep, and pigs, whereas attacks on large-sized (cow and buffalo) livestock were least frequent. Among small-sized livestock, goats are the most predated (66.92%), followed by pigs (20.30%), in all seasons. The escalating human-leopard conflict in BNP is thus a severe threat to conservation efforts as the park has already invested a substantial amount of money (approx. USD 80,000) compensating for livestock lost in leopard attacks over the last two decades. Improving habitat conditions to reduce competition inside the park, developing an insurance scheme for livestock and humans, providing support for upgraded sheds, and the development of practical and feasible strategies that focus on specific animals and clusters of the national park are needed to reduce conflicts to maintain the coexistence between wildlife and human beings.
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Protected areas (PAs) transform over time due to natural and anthropogenic processes, resulting in the loss of biodiversity and ecosystem services. As current and projected climatic trends are poised to pressurize the sustainability of PAs, analyses of the existing perturbations are crucial for providing valuable insights that will facilitate conservation management. In this study, land cover change, landscape characteristics, and spatiotemporal patterns of the vegetation intensity in the Kasungu National Park (area = 2445.10 km2) in Malawi were assessed using Landsat data (1997, 2008 and 2018) in a Fuzzy K-Means unsupervised classification. The findings reveal that a 21.12% forest cover loss occurred from 1997 to 2018: an average annual loss of 1.09%. Transition analyses of the land cover changes revealed that forest to shrubs conversion was the main form of land cover transition, while conversions from shrubs (3.51%) and bare land (3.48%) to forest over the two decades were comparatively lower, signifying a very low rate of forest regeneration. The remaining forest cover in the park was aggregated in a small land area with dissimilar landscape characteristics. Vegetation intensity and vigor were lower mainly in the eastern part of the park in 2018. The findings have implications for conservation management in the context of climate change and the growing demand for ecosystem services in forest-dependent localities.
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The causes of land degradation in the African drylands have been shown to vary. Some researchers consider climate to be the major contributor to degradation, with anthropogenic factors playing a minor role. Others reverse the significance of these two factors. A third group attributes land degradation to climate and anthropogenic factors equally. This study was undertaken to establish the factors influencing land degradation in a semi-arid environment in southeastern Kenya and the rate of change in vegetation types for a period of 35 years (1973–2007). The reduction in grassland cover was used as an indicator of land degradation. Causes of land degradation were determined by a multiple regression analysis. A log-linear regression analysis was used to establish the rate of vegetation change. The multiple and log-linear regression analyses showed: (1) woody vegetation, livestock population and cultivated area to be the main contributors of reduction in grassland cover in the area, and (2) an increase in undesirable woody species, livestock population and cultivated area had a significant (P<0.05) negative effect on grassland vegetation. Increased human population, low amounts of rainfall and drought showed no significant negative effect on grassland vegetation cover. In conclusion, human and livestock population growth and increased agricultural land have contributed to intensive crop cultivation and overgrazing in the semi-arid lands. This overuse of the semi-arid rangelands has worsened the deterioration of the natural grassland vegetation.
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Human populations continue to increase and transform Earth's ecosystems. For large carnivores, human development reduces habitat abundance, alters predator–prey dynamics, and increases the risk of mortality, which may threaten the viability of many populations. To investigate how the cougar (Puma concolor) responds to a gradient of human development in four areas in Washington, USA, we used utilization distributions, county tax parcel data, Weibull modeling analysis, and multiple comparison techniques. Cougars used wildland areas the majority of the time (79% ± 2%, n = 112 cougars), with use decreasing as housing densities increased. When present in human-developed areas in eastern Washington, 99% of the habitat that cougars used had housing densities ≤76.5 residences/km2, which was <846.0 residences/km2 observed in western Washington (P < 0.01). Cougars used areas in western Washington with greater housing density likely because of the clustered nature of housing developments, the connectivity with greenbelts and forested corridors, and security cover of dense maritime vegetation. Our findings suggest a consistent, albeit nuanced response by cougars to human development that may be used by wildlife managers, landscape planners, and environmental educators to guide and enhance their efforts to minimize the impacts of human development on cougars and reduce the potential for conflicts with people. Our model may also provide guidance for thresholds of human development for other adaptable large carnivores.
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Large carnivores’ fear of the human ‘super predator’ has the potential to alter their feeding behaviour and result in human-induced trophic cascades. However, it has yet to be experimentally tested if large carnivores perceive humans as predators and react strongly enough to have cascading effects on their prey. We conducted a predator playback experiment exposing pumas to predator (human) and non-predator control (frog) sounds at puma feeding sites to measure immediate fear responses to humans and the subsequent impacts on feeding. We found that pumas fled more frequently, took longer to return, and reduced their overall feeding time by more than half in response to hearing the human ‘super predator’. Combined with our previous work showing higher kill rates of deer in more urbanized landscapes, this study reveals that fear is the mechanism driving an ecological cascade from humans to increased puma predation on deer. By demonstrating that the fear of humans can cause a strong reduction in feeding by pumas, our results support that non-consumptive forms of human disturbance may alter the ecological role of large carnivores.
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Reports of livestock depredation by large predators were systematically collected at three study sites in northwestern Zimbabwe from 2008–2013. We recorded 1,527 incidents (2,039 animals killed and 306 injured). Lions (Panthera leo) and spotted hyaenas (Crocuta crocuta) were mostly responsible, and cattle and donkeys most frequently attacked. Patterns of predation were variable among study sites. Nevertheless, some overall patterns were apparent. Predators selected livestock close to the size of their preferred wild prey, suggesting behaviours evolved to optimise foraging success may determine the domestic species primarily preyed upon. Most attacks occurred when livestock were roaming outside and away from their 'home' protective enclosures at night. Hyaena attacks were largely nocturnal; lions and leopards (Panthera pardus) were more flexible, with attacks occurring by day and at night. Livestock fitted with bells suffered a disproportionate number of attacks; the sound of bells appears to have conditioned predators to associate the sound with foraging opportunities. Lion and hyaena attacks on cattle were more frequent in the wet season suggesting that seasonal herding practices may result in cattle vulnerability. Only a small proportion of conflict incidents were reported to wildlife management officials with a bias towards lion predation events, potentially prejudicing conflict management policies. Predation on domestic stock involves an intricate interplay between predator behaviour and ecology on the one hand and human behaviour and husbandry practices on the other. Our data suggest that improved livestock husbandry (supervision of grazing animals, protection at night in strong enclosures) would greatly reduce livestock depredation.
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There is growing recognition that developed landscapes are important systems in which to promote ecological complexity and conservation. Yet, little is known about processes regulating these novel ecosystems, or behaviours employed by species adapting to them. We evaluated the isotopic niche of an apex carnivore, the cougar (Puma concolor), over broad spatiotemporal scales and in a region characterized by rapid landscape change. We detected a shift in resource use, from near complete specialization on native herbivores in wildlands to greater use of exotic and invasive species by cougars in contemporary urban interfaces. We show that 25 years ago, cougars inhabiting these same urban interfaces possessed diets that were intermediate. Thus, niche expansion followed human expansion over both time and space, indicating that an important top predator is interacting with prey in novel ways. Thus, though human-dominated landscapes can provide sufficient resources for apex carnivores, they do not necessarily preserve their ecological relationships.
Co‐occurrence with humans presents substantial risks for large carnivores, yet human‐dominated landscapes are increasingly crucial to carnivore conservation as human land use continues to encroach on wildlife habitat. Flexibility in large carnivore behavior may be a primary factor mediating coexistence with people, allowing carnivores to calibrate their activity and habitat use to the perceived level of human risk. However, our understanding of how large carnivores adjust the timing and location of behaviors in response to variations in human activity across the landscape remains limited, impacting our ability to identify important habitat for populations outside of protected areas. Here we examine whether African lions (Panthera leo) modify their behavior and habitat use in response to risk of a human encounter, and whether behavior‐specific habitat selection allows lions to access feeding opportunities in a human‐dominated landscape in Kenya. We determined fine‐scale behavioral states for lions using high‐resolution GPS and accelerometer data, and then investigated behavior‐specific habitat selection at multiple temporal and spatial scales (ranging from fifteen minutes to twelve hours and from approximately two hundred meters to several kilometers). We found that lions exhibit substantial differences in habitat selection with respect to humans based on behavioral state and time of day. During the day, when risk of human encounter is highest, lions avoided areas of high human use when resting, meandering, and feeding. However, lions specifically selected for habitat near people when feeding at night. Flexible habitat use by lions thus permits access to prey, which appear to concentrate in areas near humans. The importance of habitat near people for feeding was only apparent when analyses explicitly accounted for lion behavioral state and spatiotemporal scale, highlighting the necessity of incorporating such information when investigating human impacts on large carnivore habitat use. Our results support the contention that behavior‐specific habitat selection promotes carnivore persistence in human‐dominated landscapes, demonstrating the importance of considering not just whether but how large carnivores use habitat near humans when managing vulnerable populations. This article is protected by copyright. All rights reserved.
Occupancy Estimation and Modeling: Inferring Patterns and Dynamics of Species Occurrence, Second Edition, provides a synthesis of model-based approaches for analyzing presence-absence data, allowing for imperfect detection. Beginning from the relatively simple case of estimating the proportion of area or sampling units occupied at the time of surveying, the authors describe a wide variety of extensions that have been developed since the early 2000s. This provides an improved insight about species and community ecology, including, detection heterogeneity; correlated detections; spatial autocorrelation; multiple states or classes of occupancy; changes in occupancy over time; species co-occurrence; community-level modeling, and more. Occupancy Estimation and Modeling: Inferring Patterns and Dynamics of Species Occurrence, Second Edition has been greatly expanded and detail is provided regarding the estimation methods and examples of their application are given. Important study design recommendations are also covered to give a well rounded view of modeling.
The loss of apex consumers (large mammals at the top of their food chain) is a major driver of global change [1]. Yet, research on the two main apex consumer guilds, large carnivores [2] and megaherbivores [3], has developed independently, overlooking any potential interactions. Large carnivores provoke behavioral responses in prey [1, 4], driving prey to distribute themselves within a "landscape of fear" [5] and intensify their impacts on lower trophic levels in low-risk areas [6], where they may concentrate nutrients through localized dung deposition [7, 8]. We suggest, however, that megaherbivores modify carnivore-induced trophic cascades. Megaherbivores (>1,000 kg [9]) are largely invulnerable to predation and should respond less to the landscape of fear, thereby counteracting the effects of fear-triggered trophic cascades. By experimentally clearing plots to increase visibility and reduce predation risk, we tested the collective role of both apex consumer guilds in influencing nutrient dynamics in African savanna. We evaluated whether megaherbivores could counteract a behaviorally mediated trophic cascade by redistributing nutrients that accumulate through fear-driven prey aggregations. Our experiment showed that mesoherbivores concentrated fecal nutrients in more open habitat, but that megaherbivores moved nutrients against this fear-driven nutrient accumulation by feeding within the open habitat, yet defecating more evenly across the risk gradient. This work adds to the growing recognition of functional losses that are likely to have accompanied megafaunal extinctions by contributing empirical evidence from one of the last systems with a functionally complete megaherbivore assemblage. Our results suggest that carnivore-induced trophic cascades work differently in a world of giants.
Leopard (Panthera pardus) populations across Africa are increasingly exposed to high levels of anthropogenic disturbance, and information on habitat use responses of leopards in human-disturbed landscapes can help inform status assessments and guide conservation interventions. Unfortunately, however, few studies have investigated leopard ecology in human-disturbed landscapes, particularly in Africa. We employed camera-trapping and occupancy modelling to provide inferences on leopard habitat use in a National Park in Mozambique impacted by subsistence farming and bushmeat poaching. Replicated detection/non-detection occupancy surveys were used to estimate site use by leopards in a representative area of the park, and to investigate relative impacts of environmental, conspecific and anthropogenic factors on leopard occurrence. The proportion of sites used by leopards was estimated at 0.814 (SE = 0.093), which is approximately twice the occupancy previously reported for lion (44%) and cheetah (40%) in the same area. Leopard presence was not strongly predicted by any of the covariates, indicating there were no strong limiting factors. While leopards generally avoided human settlements and were positively predicted by prey, results suggest that there was sufficient prey and space for the species to use most available habitats. The greatest contributing factor to leopard habitat use was a positive correlation with bushmeat poachers and lions. It is possible that these other predators provide a more accurate indicator of prey availability than our single-species indicator based on camera trap data. This study provides important novel information on habitat use by leopards in a system disturbed by rural human subsistence activities in Africa.