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Spatial segregation between Iberian lynx and other carnivores

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Abstract and Figures

Rabbits represent the bulk of its diet as for many other Iberian predators. This study addresses how the presence of the Iberian lynx affects the spatial distribution of the mesocarnivore community at landscape scale in the Sierra de Andújar. We studied mesocarnivore presence by sampling at 230 camera trapping stations, located in areas with and without lynx. We used a x2–test to compare the proportion of stations in which each species of carnivore were recorded in the zones with and without lynx. The proportion of camera trapping stations in which red fox (Vulpes vulpes), Egyptian mongoose (Herpestes ichneumon), beech marten (Martes foina), wildcat (Felis sylvestris) and common genet (Genetta genetta) were detected was significantly lower in the area where lynx were present than in the area where it was absent. No significant differences between the two types of areas were found for badgers (Meles meles). Our results highlight the role of the lynx as apex predators and the benefits that the recovery of Iberian lynx populations would entail in terms of trophic interactions and restored disrupted ecosystems processes.Key words: Intraguild competition, Carnivores, Phototrapping, Apex preda
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Animal Biodiversity and Conservation 42.2 (2019)
© 2019 Museu de Ciències Naturals de Barcelona
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ISSN: 1578–665 X
eISSN: 2014–928 X
Garrote, G., Pérez de Ayala, R., 2019. Spatial segregation between Iberian lynx and other carnivores. Animal
Biodiversity and Conservation, 42.2: 347–354, Doi:
Spatial segregation between Iberian lynx and other carnivores. The Iberian lynx (Lynx pardinus) is a specialist
predator. Rabbits represent the bulk of its diet as for many other Iberian predators. This study addresses how
the presence of the Iberian lynx affects the spatial distribution of the mesocarnivore community at landscape
scale in the Sierra de Andújar. We studied mesocarnivore presence by sampling at 230 camera trapping
stations, located in areas with and without lynx. We used a x2–test to compare the proportion of stations in
which each species of carnivore were recorded in the zones with and without lynx. The proportion of camera
trapping stations in which red fox (Vulpes vulpes), Egyptian mongoose (Herpestes ichneumon), beech marten
(Martes foina), wildcat (Felis sylvestris) and common genet (Genetta genetta) were detected was signicantly
lower in the area where lynx were present than in the area where it was absent. No signicant differences
between the two types of areas were found for badgers (Meles meles). Our results highlight the role of the
lynx as apex predators and the benets that the recovery of Iberian lynx populations would entail in terms of
trophic interactions and restored disrupted ecosystems processes.
Key words: Intraguild competition, Carnivores, Phototrapping, Apex predator
Segregación espacial entre el lince ibérico y otros carnívoros. El lince ibérico (Lynx pardinus) es un depreda-
dor especialista. El conejo constituye el grueso de su dieta, al igual que la de otros depredadores ibéricos.
Este estudio analiza cómo la presencia del lince ibérico afecta a la distribución espacial de la comunidad de
mesocarnívoros a escala de paisaje en la sierra de Andújar. Se estudió la presencia de mesocarnívoros me-
diante 230 cámaras de fototrampeo, instaladas en zonas con y sin presencia de lince. Se utilizó la prueba de
la x2 para comparar la proporción de cámaras en las que se detectó cada una de las especies de carnívoros
en las zonas con y sin lince. La proporción de cámaras que detectaron zorros (Vulpes vulpes), meloncillos
(Herpestes ichneumon), garduñas (Martes foina), gatos monteses (Felis sylvestris) y ginetas (Genetta genetta)
fue signicativamente menor en las zonas con presencia de lince que en las zonas donde este estaba ausente.
No se encontraron diferencias signicativas en cuanto a la presencia de tejones (Meles meles) entre ambos
tipos de zona. Nuestros resultados ponen de relieve la importancia del lince como depredador apical y los
benecios que podría reportar la recuperación de las poblaciones de lince ibérico en lo que concierne a las
interacciones trócas y el restablecimiento de los procesos ecosistémicos interrumpidos.
Palabras clave: Competencia intragremial, Carnívoros, Fototrampeo, Depredador apical
Received: 08 XI 18; Conditional acceptance: 05 II 19; Final acceptance: 16 V 19
Germán Garrote, Instituto de Biología de la Conservación (IBiCo), c/ Nebli 13, 28232, Madrid, España (Spain).–
R. Pérez de Ayala, WWF/España, Gran Vía de San Francisco 8–D, 28005 Madrid, España (Spain).
Corresponding author: G. Garrote. E–mail:
Spatial segregation between
Iberian lynx and other carnivores
G. Garrote, R. Pérez de Ayala
348 Garrote and Pérez de Ayala
Direct interactions between predators and other species
can have indirect consequences further down the food
web via trophic cascades (Ripple et al., 2016). Large
carnivores play a key role in terrestrial ecosystems
when they exert an inuence on herbivores and so
indirectly prevent overgrazing (McShea, 2005). They
can also inuence carnivore communities via intraguild
interactions (Ritchie and Johnson, 2009) and indirectly
prevent excessive predation on prey species by meso-
carnivores (Elmhagen et al., 2010). This top–down
cascade can inuence ecosystem structures and
biodiversity at both local and larger scales (Terborgh,
2001; Elmhagen et al., 2010). If healthy populations
of top predators are to be maintained within ecosys-
tems, these ecosystems should also contain healthy
communities and populations of the many species that
perform ecosystem services at lower trophic levels
(Dobson et al., 2006; Haswell et al., 2017). However,
the functional roles of top predators cannot be fully
appreciated in isolation from bottom–up processes
because the effects of nutrients, productivity (Pace et
al., 1999) and anthropogenic habitat may bring about
change (Litvaitis and Villafuerte, 1996; Estes, 1998;
Elmhagen and Rushton, 2007).
Competitive intraguild interactions have been propo-
sed as highly important organizing mechanisms since,
due to similarities in ecological niches, they limit the
number of species that can be packed into an assem-
blage (Jaksic and Marone, 2007). Similar ecological
preferences increase the risk of competition, whereas
mechanisms such as resource partitioning, temporal
or spatial avoidance strategies (Voigt and Earle, 1983;
Johnson and Franklin, 1994; Kozlowski et al., 2008),
or alternative foraging strategies (Husseman et al.,
2003) facilitate coexistence. Interference interactions,
harassment and injury caused by larger carnivores pose
a risk to smaller mesopredators (Linnell and Strand,
2000; Haswell et al., 2018). Furthermore, as a result
of interference competition, subordinate species are
frequently restricted to suboptimal habitats (Tannerfeldt
et al., 2002; Macdonald et al., 2004; Mitchell and Banks,
2005), which can have important implications for the
demography and distribution of the species involved
(Thompson, 1988; Holt and Polis, 1997; Atwood and
Gese, 2008).
The Iberian lynx (Lynx pardinus) is the top pre-
dator of the terrestrial vertebrate community in the
Mediterranean ecosystem (Valverde, 1963). Listed as
Endangered by the IUCN (Rodríguez and Calzada,
2015), the species reached its all–time minimum in the
rst years of the twenty–rst century, when only 100
individuals in just two isolated populations –Andújar–
Cardeña and Doñana– were known to exist (Guzmán
et al., 2004; Simón et al., 2012). Since then, however,
the Iberian lynx has undergone a signicant increase
in population size and range due to the measures
implemented as part of conservation projects for
the species (Simón et al., 2012), which include the
creation of new populations through reintroduction.
The Iberian lynx is a specialist predator. Rabbits
represent the bulk of its diet in a similar manner to
that of many other Iberian predators (Cabezas–Díaz
et al., 2011), possibly leading to interference or
food competition. Previous studies of the relation-
ships between Iberian lynx and other carnivores
performed in Doñana have found that the Egyp-
tian mongoose (Herpestes ichneumon) and genet
(Genetta genetta) avoid lynx, while the Eurasian
badger (Meles meles) is apparently indifferent to
its presence. Although foxes (Vulpes vulpes) and
lynx exhibit temporal segregation in their use of
habitat (Fedriani et al., 1999), their spatial rela-
tionship remains unclear (Palomares et al., 1996).
The relationship between wildcat (Felix sylvestris)
and lynx has not been studied.
This study addresses how the presence of the
Iberian lynx affects the spatial distribution of the
mesocarnivore community at a landscape scale in the
Sierra de Andújar. We studied the spatial distribution
of several species of mesocarnivores in areas where
the lynx is absent and where it is present, taking into
account the abundance of rabbits.
Material and methods
Study area
The study area lies in the eastern Sierra Morena (SE
Spain; g. 1) and consists of a mountainous area
with an altitudinal range of 200–1,500 m covered by
well–preserved Mediterranean forests (Quercus ilex,
Q. faginea and Q. suber) and scrublands (Quercus
coccifera, Pistacia lentiscus, Arbutus unedo, Phillyrea
angustifolia and Myrtus communis). The area is ma-
naged for big–game hunting and has high densities
of red deer (Cervus elaphus) and wild boar (Sus
scrofa). It is partially protected by the Parque Natu-
ral Sierra de Andújar. During the study period, the
Andújar–Cardeña Iberian lynx population consisted
of 60–110 individuals, distributed over an area of
15,000 ha (Guzmán et al., 2004).
Camera trapping survey
The spatial distribution of the carnivore community was
estimated by sequential camera trapping surveys per-
formed in December 1999–February 2000, November
2000–February 2001 and November 2001–February
2002. We used camera trapping data from the annual
national Iberian lynx survey (Guzmán et al., 2004),
which covers 85 % of the area potentially used by the
Iberian lynx.
We divided the study area into 12 survey blocks,
each of which were surveyed by camera trapping for
periods of two months. Once one block was nis-
hed, cameras were moved to the next survey block.
We surveyed an almost continuous surface area of
7,800 ha using a total of 230 camera trapping stations
(1999/2000: n = 28; 2000/2001: n = 168; 2001/2002:
n = 39). In all, 115 out of 230 stations were located
in areas in which the lynx are present, as dened by
Guzman et al. (2004), and the other 115 stations were
placed in areas without lynx (g. 1).
Animal Biodiversity and Conservation 42.2 (2019) 349
Fig. 1. Study area map. Camera trap stations located in areas with and without lynx, and stations in
which each species of carnivore was recorded.
Fig. 1. Mapa de la zona de estudio donde se representa la ubicación de las estaciones de fototrampeo en
zonas con y sin lince, así como las estaciones donde se detectó cada una de las especies de carnívoros.
Camera trap
2002 Iberian lynx
distribution area
2 0 2 4 k m
Felis sylvestris Meles meles
Vulpes vulpes Martes foina
Genetta genetta Herpestes ichneumon
350 Garrote and Pérez de Ayala
We used 212 35–mm Canon Prima© classic photo
lm cameras with data registers and automatic ashes.
The cameras were modied to allow activation via an
external 25 × 25 cm pressure plate, positioned at a
distance of 170 cm that was triggered when stepped
on by an animal (Garrote et al., 2011). The cameras
were placed in a small wooden box on pillars 30 cm
above ground level. Urine from captive Iberian lynx,
placed on an inert adjacent support, 50 cm above
ground level and the pressure plate, was used as a
lure. Lynx urine has been reported to be an excellent
attractant for all carnivore species (Garrote et al., 2011;
Monterroso et al., 2016). This attractant was replaced
every 3–6 days. The distance between camera traps
was 400–800 m. Camera–trap locations were located
along suspected lynx travel routes (Garrote et al., 2012)
such as roads or paths, chosen to maximize capture
probabilities (Karanth and Nichols, 1998). Each camera
was continuously active throughout the entire survey
period for each block (two months).
To describe the species distribution in the area, we
calculated occupancy as the proportion of stations at
which a species was detected in relation to the total
number of stations (Sogbohossou et al., 2018).
Rabbit abundance and habitat variables
Rabbit abundances were estimated for each survey
block by on–foot constant–speed itineraries lasting
three hours. Rabbit latrines were counted every 15',
and these counts were taken as the survey unit for
the statistical analysis. Indirect surveys were carried
out at the same time of the year (end of spring,
when rabbit populations peak) under similar weather
conditions. Every 15' we estimated, in a 25 m radius
plot, the percentage of land surface covered by the
following habitat categories: trees, scrubland lower
than 50 cm in height, scrubland higher than 50 cm
in height, pastureland and rocks. The percentage of
covered land was divided into four categories scored
as follows: 1 (0–25 %), 2 (> 25–50 %), 3 (> 50–75 %)
and 4 (> 75).
Statistical analysis
We compared the mean values for rabbit abundance
and for each habitat category obtained in the areas
with and without lynx using a Mann–Whitney U–test.
We used a x2–test to compare the proportion of
stations in the zones with and without lynx in which
each species of mesocarnivore was present. The
carnivores with lower capture rates were grouped
together to perform statistical analysis (minimum ve
expected records).
The following carnivores were detected in this study:
(Lynx pardinus, 9–15.9 kg), Eurasian badger (Meles
meles), red fox (Vulpes vulpes), Egyptian mongoo-
se (Herpestes ichneumon), beech marten (Martes
foina), wildcat (Felis sylvestris), and common genet
(Genetta genetta). The proportion of camera trapping
stations in which the fox and wildcat were detected
was signicantly lower in the area with lynx than in
the area without lynx (table 1; g. 1); no signicant
differences were found for the presence of the bad-
ger between both areas. Genet, beech marten and
Egyptian mongoose were grouped together to perform
the statistical analysis. The presence of this group of
mesocarnivores was found to be signicantly lower
in the areas where lynx were present.
No signicant difference was found between zones
with and without lynx for the habitat variables (ta-
ble 2). As expected, rabbit abundance in areas with
lynx was signicantly higher than that in lynx–free
areas since lynx distribution is dependent on rabbit
abundance (table 2).
With the exception of the badger, the presence of the
Iberian lynx determines the distribution at the lands-
cape scale of the mesocarnivores community in the
study areas. No signicant habitat differences were
found between areas with and without lynx, while the
highest rabbit abundances were detected in areas with
lynx. As mentioned above, Iberian mesocarnivores
preferably select rabbits as prey (Cabezas–Díaz et
al., 2011). The most probable explanation for the ob-
served distribution of mesocarnivores at a landscape
scale is the interference competition between species
in which the lynx is the dominant species.
This is the rst study to address a relationship
between the Iberian lynx and wildcat, the only two
sympatric wild felids present in the Iberian peninsula.
Competition becomes greater as eco–morphological
similarities or phylogenetic proximity between compet-
ing species increase (Cruz et al., 2018), and generally
the larger dominant species exclude smaller or sub-
ordinate species from their territories by interference
competition. Therefore, as expected, the larger Iberian
lynx exerts strong interference competition on the
smaller wildcat. This leads to fewer wildcats in those
areas where lynx are present. Similar relationships
of dominance have been described for other species
of felines, such as the ocelot (Leopardus pardalis),
which acts as a dominant carnivore over other smaller
sympatric cats such as margay (Leopardus wiedii) and
jaguarundi (Puma yagouaroundi) and so inuences
their ecological parameters (de Oliveira et al., 2010;
Cruz et al., 2018).
Previous studies have shown a high overlap in the
diets, activity levels, habitat use and home range in
radio–tracked foxes and lynx (Fedriani et al., 1999).
Although it has been suggested that foxes mitigate
lynx predation by modifying their spatial behaviour
at home range level, no spatial segregation in these
species has ever been found. Using a landscape
approach, the present study demonstrates signicant
spatial segregation between foxes and lynx. These
differences with previous work might be attributable
to scale since certain studies have concluded that
approaches at different scales can generate different
Animal Biodiversity and Conservation 42.2 (2019) 351
conclusions regarding interspecic interactions bet-
ween species (e.g. (Tannerfeldt et al., 2002) for the
Arctic red fox (Cruz et al., 2018). Previous studies
(Palomares et al., 1996; Fedriani et al., 1999) have
covered smaller areas than our study, which was
performed at a much greater landscape scale. On the
other hand, the relative densities of the mesocarni-
vores and their prey may also inuence interactions
(Creel, 2001; Berger and Gese, 2007). However,
although no information is available for fox densities
to compare these two study areas, the density of Ma-
tasgordas rabbit population (8 rabbits/ha; Villafuerte
et al., 1997) is greater than that of Andújar (Simón
et al., 2012). In areas or during periods of lower prey
abundance, competition may play a more important
role and interspecic interactions may change, re-
sulting in increased interference competition (Creel,
2001). Lower prey densities can result in lower lynx
tolerance toward foxes and, consequently, greater
interference competition. Similar conclusions were
reached by (Gese et al., 1996) in Yellowstone National
Park, where coyotes tolerate red foxes during high
prey years but not at other times.
Although data regarding the presence of the smaller
mesocarnivores (Mongoose, martens and genets) are
scarce, our results concur with previously reports from
Doñana, where mongoose and genets avoid areas
where lynx are present.
Iberian lynx and badgers seem to be particularly
well predisposed to coexist (Palomares et al., 1996;
Fedriani et al., 1999), and our results suggest that
there is a complete spatial overlap between the
species. Kleiman and Eisenberg (1973) suggest that
this coexistence occurs as a result of a separation in
their ecological niches, which is likely a consequen-
ce of evolution of different social systems. Similar
interactions have been described between Eurasian
lynx and wolves in Białowieza Forest (Schmidt, 2008)
and between lynx and wolverine in northern Sweden
(Schmidt, 2008). The Iberian lynx is a crepuscular
species that preys mainly on rabbits (Fedriani et al.,
1999), whereas badgers are much more nocturnal
and are generalists with the capacity to survive on a
Table 1. Total number of camera stations, positive stations for each species in zones with/without
lynx, and positive stations per species. Genet, beech marten and Egyptian mongoose are grouped in
'others'. x2 results are shown.
Tabla 1. Número total de estaciones de fototrampeo, número de estaciones positivas para cada especie en
las zonas con y sin lince y estaciones totales positivas para cada especie. Las ginetas, las garduñas y los
meloncillos están agrupados en la categoría "others" (otras). Se muestran los resultados de las pruebas de la x2.
Total Badger Fox Wildcat Others
With lynx 115 20 10 6 2
Without lynx 115 26 53 29 17
Total 230 43 63 35 19
p 0.5 < 0.0001 0.0038 0.016
Table 2. Mann–Whitney U–test results for the
variables of habitat and rabbit abundance.
Tabla 2. Resultados de las pruebas U de
MannWhitney para las variables del hábitat y
la abundancia de conejos.
Z p–level
Pasture 0.64 0.52
Scrub < 50cm 0 1
Scrub > 50 cm 0.96 0.33
Tree 0.48 0.63
Rocks 1.28 0.2
Rabbit 2.08 0.03
greater diversity of resources (Roper, 1994; Neal and
Cheeseman, 1996; Revilla and Palomares, 2002).
The food available for badgers in Mediterranean ha-
bitats varies greatly and badgers respond by shifting
their diets accordingly between prey items (Virgós et
al., 2004). However, niche differences alone cannot
completely explain this coexistence. Foxes are even
more adaptable than badgers and could potentially
develop resource partitioning, temporal avoidance
strategies (Voigt and Earle, 1983; Johnson and
Franklin, 1994; Kozlowski et al., 2008), or different
foraging strategies (Husseman et al., 2003) to facili-
tate coexistence. However, fox distribution is clearly
inuenced by the presence of lynx while badger
distribution is not. The outcome of direct encounters
between lynx and badgers is unknown but probably
involves a risk of injury for both species. Therefore,
the observed sympatry between Iberian lynx and
badger is probably facilitated by a combination of
both factors –the avoidance of injury and different
foraging strategies.
352 Garrote and Pérez de Ayala
As a result of being a trophic specialist on rabbits,
the abundance of its staple prey determines the lynx’s
basic demographic parameters (Monterroso et al.,
2016) and distribution (Guzmán et al., 2004), which
thus implies that there is bottom–up control over Iberian
lynx dynamics. Likewise, the presence or absence of
the Iberian lynx, which is determined by rabbit abun-
dance, affects the dynamics of subordinate carnivore
species via a top–down control effect. The foraging
theory suggests that animals adjust their behaviour
accordingly to optimize foraging efciency and overall
tness, and trade–off harvesting rates with tness costs
(Haswell et al., 2018). In the absence of Iberian lynx,
sympatric mesocarnivores should ideally be distribu-
ted on the basis of habitat quality and preferred food
availability (Van Der Meer and Ens, 1997; Roemer et
al., 2009). The presence of the lynx forces smaller
species to invest in antipredator behavioural strate-
gies (Lima, 1998; Haswell et al., 2017) that can have
negative consequences. For example, their access to
high–quality foraging areas can be restricted (Ritchie
and Johnson, 2009), which forces them to seek an
alternative diet, adopt their life cycles to those of their
new prey items, and adjust their feeding behaviour
(Durant, 2000; Hayward and Slotow, 2009; Wikenros
et al., 2014). This in turn can affect the size of the
home range, increase travel costs or lead to shifts in
habitat use (Caro and Stoner, 2003). The tness costs
of these antipredator responses could affect survival
and reproduction, thereby ultimately having an impact
on population dynamics (Creel and Christianson,
2008). On the other hand, a fall in lynx numbers is
expected following rabbit declines, which will lead to a
lessening of the top–down control on mesocarnivores
numbers (Estes et al., 2011; Monterroso et al., 2016).
Conservation implications
Numerous studies have drawn attention to the impor-
tance of apex predators in suppressing populations
of smaller predators (mesopredators) and thus their
roles in moderating the impact of predation on sma-
ller prey species (Crooks and Soulé, 1999; Johnson
et al., 2007; Berger et al., 2008). The recovery and
re–establishment of apex predator populations con-
tribute not only to their conservation but also benet
biodiversity conservation via a relaxing of the impact
of mesopredators on their prey (Ritchie and Johnson,
2009). This is positive for the restoration of disrupted
ecosystem processes (Estes et al., 2011; Ritchie et
al., 2012), particularly in terms of trophic interactions
(Monterroso et al., 2016) but also for economic and
social reasons (ecosystem services). Some areas in
rural Spain have high rabbit densities and suitable
habitat for the lynx. Most such areas are occupied
by private, intensively managed, small–game hun-
ting areas (rabbit and partridge; Delibes–Mateos et
al., 2009). In these hunting estates strong predator
control is traditional and still persists nowadays, both
legally (leg–hold traps and snares when authorised
under certain exceptional circumstances) and illegally
(Villafuerte et al., 2000; Virgós and Travaini, 2005).
Despite the possible negative effect on non–target
species, this practice requires important time and
monetary expenditure, although the desired results
are not always achieved (Harding et al., 2001). Lynx
are viewed negatively by many hunters in the Iberian
Peninsula since, as a trophic specialist that preys on
rabbits, it competes for this highly important small–
game species. Nevertheless, the Iberian lynx presen-
ce could be an effective, natural and inexpensive tool
for predator control since it suppresses populations
of smaller predators and thereby mitigates the impact
that these mesopredators will have on game species
(Palomares et al., 1995). This is a key argument for
changing game managers’ opinions and for ensuring a
favourable response to any lynx reintroduction project
in its past range from where, ironically, it was eradi-
cated by indiscriminate predator control (Gil–Sánchez
and McCain, 2011).
This study was supported by DGCONA–MIMAM
project 'Censo–Diagnóstico de las poblaciones de
Lince Ibérico en España', and by IBiCO/WWF Spain/
Fundación Barcelona Zoo project 'Competencia inte-
respecíca y coexistencia entre el lince ibérico (Lynx
pardinus) y otros carnívoros'. We wish to express our
gratitude to Nicolas Guzmán, Paco García, Aquilino
Duque and Concha Iglesias who carried out eldwork
with us. We also thank the Organismo Autónomo de
Parques Nacionales (Lugar nuevo), Parque Natural
de la Sierra de Andújar, TRAGSA, Fundación CBD–
Habitat, WWF/España, EGMASA, CMA Junta de
Andalucía. We thank Jose Luis Tellería and Guillermo
López for their constructive comments.
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... The large dataset included in the present study strongly validates the competitive exclusion between the lynx and the fox already described in Garrote and Ayala (2019) and Jiménez et al. (2019). However, a reintroduction effect on another predator s abundance and spatial and time use was not reported in Garrote and Ayala (2019), since the lynx was never completely absent from the study area. ...
... The large dataset included in the present study strongly validates the competitive exclusion between the lynx and the fox already described in Garrote and Ayala (2019) and Jiménez et al. (2019). However, a reintroduction effect on another predator s abundance and spatial and time use was not reported in Garrote and Ayala (2019), since the lynx was never completely absent from the study area. Nonetheless, spatial segregation between lynxes and foxes was also observed in Garrote and Ayala (2019). ...
... However, a reintroduction effect on another predator s abundance and spatial and time use was not reported in Garrote and Ayala (2019), since the lynx was never completely absent from the study area. Nonetheless, spatial segregation between lynxes and foxes was also observed in Garrote and Ayala (2019). As for Jiménez et al. (2019), the authors did analyse the effect of lynx reintroduction on foxes and also small game abundances. ...
... The large dataset included in the present study strongly validates the competitive exclusion between the lynx and the fox already described in Garrote and Ayala (2019) and Jiménez et al. (2019). However, a reintroduction effect on another predator s abundance and spatial and time use was not reported in Garrote and Ayala (2019), since the lynx was never completely absent from the study area. ...
... The large dataset included in the present study strongly validates the competitive exclusion between the lynx and the fox already described in Garrote and Ayala (2019) and Jiménez et al. (2019). However, a reintroduction effect on another predator s abundance and spatial and time use was not reported in Garrote and Ayala (2019), since the lynx was never completely absent from the study area. Nonetheless, spatial segregation between lynxes and foxes was also observed in Garrote and Ayala (2019). ...
... However, a reintroduction effect on another predator s abundance and spatial and time use was not reported in Garrote and Ayala (2019), since the lynx was never completely absent from the study area. Nonetheless, spatial segregation between lynxes and foxes was also observed in Garrote and Ayala (2019). As for Jiménez et al. (2019), the authors did analyse the effect of lynx reintroduction on foxes and also small game abundances. ...
... Soto-Navarro 2013), they are less common in Coto del Rey, the only zone where lynxes are present continuously in the National Park and a high-density rabbit population exists. Moreover, the intolerance of the presence of the Iberian lynx has been established as one of the main reasons why mesocarnivores avoid areas occupied by lynx, which leads to spatial segregation (Palomares et al. 1996;Palomares and Caro 1999;Alonso and de Ayala 2019). The red fox exhibits fine-scale habitat and temporal avoidance of lynxes, which allows them to co-exist to some extent (Soto-Navarro 2013;Soto and Palomares 2015), even though lynx predate on foxes as well as on other mesocarnivore species (Palomares and Caro 1999). ...
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Long-term studies of community composition and relative abundance are key tools in wildlife management and biodiversity conservation. However, few studies of this kind are available for Mediterranean carnivores, especially in the Iberian Peninsula , a hotspot of mammal biodiversity in Europe. We used 15 years of carnivore monitoring data from the Doñana National Park, one of the most representative areas for carnivores in Iberia, to obtain population trends for the main Mediterranean carnivore species. They were positive for red fox, stable for badger and Egyptian mongoose, and negative for common genet and Iberian lynx. The importance of long-term datasets and the implications of the results for the studied species at global level are discussed, above all for species whose population trends are less well known. This is the case of the Egyptian mongoose, for which we present novel information on its long-term population trend in Europe, and of the Iberian lynx, an endangered species with a clear negative trend in this well-protected area.
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This guidance reviews the methods for estimating relative abundance and density in nine large European wild carnivore species, somerepresenting relevant health concerns andprovides insights on how to obtain reliable estimations by using those methods. On a local scale, the appropriate method should take into accountthe characteristics of the study area, the estimated survey efforts, the expected results (i.e. a measure of true density or just an index of abundance to monitor the trend in space and time) the level of accuracy and precision, and a proper design so to obtain a correct interpretation of the data. Among all methods, the camera trapping (CT) methods, especially those recently developed, are the most promising for the collection of robust data and can be conducted in a wide range of species, habitats, seasons and densities with minimal adjustments. Some recently developed CT methods do not require individual recognition of the animals and are a good compromise of cost, effort and accuracy. Linear transects,particularly Kilometric Abundance Index (KAI) is applicable for monitoring large regions.A large challenge is compiling and validating abundance data at different spatial scales. Based on ENETWILD initiative, we recommend developing a permanent network and a data platform to collect and share local density estimates, so as abundance in the EU, which would enable to validate predictions for larger areas by modelling. It would allow to identify gaps in the data on wild carnivores (including the species not assessed in the present report) and to focus on these areas for improving predictions. This platform must facilitate the reporting by wildlife policy makers and relevant stakeholders, but also citizen science initiatives.Also, there is need to improve the reliability of local density estimations by developing practical research on methods able to derive densities in untested species and situations, making the application of methods easier for local teams.
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Competition theory and niche theory suggest that two morphologically similar species may coexist by reducing the overlap of at least one dimension of their ecological niche. The medium and small Neotropical felids are an interesting group of carnivore species for studying intraguild competition. Due to differences in size it is expected that the larger ocelot exert strong interference competition on the smaller felids (southern tiger cat, margay and jaguarundi); which, in turn, may exert exploitative competition among themselves. Moreover, landscape changes due to human activities may alter these interspecific interactions. We studied the habitat use and the spatial and temporal interspecific relations of the medium and small Atlantic Forest felids, in a landscape with different levels of anthropogenic impact. We estimated the detection probability, and occupancy probability of these cats and whether these parameters are affected by environmental and anthropogenic variables or by the estimated occupancy and detection probability of the ocelot. We estimated the overlap in daily activity patterns between pairs of the four species and changes in their activity in response to anthropogenic impact. We also studied the potential changes that may have occurred in the daily activity of the small felids in relation to ocelot's occupancy probability. The probability of habitat use of the small- and medium-size felids was negatively associated to the intensity of landscape use by humans. Co-occurrence models indicated that the probability of habitat use by southern tiger cats decreased with ocelot occupancy probability. This effect was higher as human disturbance increased. Moreover, the ocelot and the southern tiger cat became more nocturnal in sites with higher human access, suggesting that they may be temporally avoiding encounters with humans or dogs. Conservation of medium and small felids in the Atlantic Forest depends not only on the establishment and implementations of protected areas but also on the management of human's land uses.
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Where direct killing is rare and niche overlap low, sympatric carnivores may appear to coexist without conflict. Interference interactions, harassment and injury from larger carnivores may still pose a risk to smaller mesopredators. Foraging theory suggests that animals should adjust their behaviour accordingly to optimise foraging efficiency and overall fitness, trading off harvest rate with costs to fitness. The foraging behaviour of red foxes, Vulpes vulpes, was studied with automated cameras and a repeated measures giving-up density (GUD) experiment where olfactory risk cues were manipulated. In Plitvice Lakes National Park, Croatia, red foxes increased GUDs by 34% and quitting harvest rates by 29% in response to wolf urine. In addition to leaving more food behind, foxes also responded to wolf urine by spending less time visiting food patches each day and altering their behaviour in order to compensate for the increased risk when foraging from patches. Thus, red foxes utilised olfaction to assess risk and experienced foraging costs due to the presence of a cue from gray wolves, Canis lupus. This study identifies behavioural mechanisms which may enable competing predators to coexist, and highlights the potential for additional ecosystem service pathways arising from the behaviour of large carnivores. Given the vulnerability of large carnivores to anthropogenic disturbance, a growing human population and intensifying resource consumption, it becomes increasingly important to understand ecological processes so that land can be managed appropriately. Electronic supplementary material The online version of this article (10.1007/s00442-018-4133-3) contains supplementary material, which is available to authorized users.
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Large predators in West Africa are threatened with extinction mainly by direct and indirect effects of human activities. Within this context, intraguild competition can limit populations of some species and even play a role in extinction. In this study, we used camera trapping to assess the spatial and temporal patterns of niche partitioning between the African lion Panthera leo leo and the spotted hyena Crocuta crocuta in Pendjari Biosphere Reserve, Benin. We found that these predators are more nocturnal in the hunting zone than in the national park of the biosphere reserve. The temporal overlap between lion and hyena was high in the national park (Pianka overlap index 0.88) and low in the hunting zones (0.39). The spatial overlap was low (0.40 in the national park and 0.38 in the hunting zones). The two predators were distributed independently in the national park, but showed significant positive association (co-occurrence) in the hunting zones. We suggest that anthropogenic activities leading to depletion of predators and their prey limit lion and hyena distribution in the hunting zones to some safety areas which are strongly selected by both predators. We recommend to significantly improve conservation efforts in the hunting zones of Pendjari Biosphere Reserve and to expand research of lion-hyena intraguild relationships to improve predator survival in West Africa.
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Emergent diseases may alter the structure and functioning of ecosystems by creating new biotic interactions and modifying existing ones, producing cascading processes along trophic webs. Recently, a new variant of the rabbit haemorrhagic disease virus (RHDV2 or RHDVb) arguably caused widespread declines in a keystone prey in Mediterranean ecosystems - the European rabbit (Oryctolagus cuniculus). We quantitatively assess the impact of RHDV2 on natural rabbit populations and in two endangered apex predator populations: the Iberian lynx (Lynx pardinus) and the Spanish Imperial eagle (Aquila adalberti). We found 60–70% declines in rabbit populations, followed by decreases of 65.7% in Iberian lynx and 45.5% in Spanish Imperial eagle fecundities. A revision of the web of trophic interactions among rabbits and their dependent predators suggests that RHDV2 acts as a keystone species, and may steer Mediterranean ecosystems to management-dependent alternative states, dominated by simplified mesopredator communities. This model system stresses the importance of diseases as functional players in the dynamics of trophic webs.
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Interactions between large carnivores and other species may be responsible for impacts that are disproportionately large relative to their density. Context-dependent interactions between species are common but often poorly described. Caution must be expressed in seeing apex predators as ecological saviours because ecosystem services may not universally apply, particularly if inhibited by anthropogenic activity. This review examines how the impacts of large carnivores are affected by four major contexts (species assemblage, environmental productivity, landscape, predation risk) and the potential for human interference to affect these contexts. Humans are the most dominant landscape and resource user on the planet and our management intervention affects species composition, resource availability, demography, behaviour and interspecific trophic dynamics. Humans can impact large carnivores in much the same way these apex predators impact mesopredators and prey species - through density-mediated (consumptive) and trait/behaviourally-mediated (non-consumptive) pathways. Mesopredator and large herbivore suppression or release, intraguild competition and predation pressure may all be affected by human context. The aim of restoring ‘natural’ systems is somewhat problematic and not always pragmatic. Interspecific interactions are influenced by context, and humans are often the dominant driver in forming context. If management and conservation goals are to be achieved then it is pivotal to understand how humans influence trophic interactions and how trophic interactions are affected by context. Trade-offs and management interventions can only be implemented successfully if the intricacies of food webs are properly understood.
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European rabbits (Oryctolagus cuniculus) are the major prey in Spain for several endangered carnivores and raptors. In recent years, many different efforts have been directed at increasing rabbit populations, and thus enhancing the survival of these endangered predators. The aim of this work was to improve our understanding of the factors that influence the natural variation in rabbit populations, specially those regarding on the food abundance and quality, and to discuss the efforts actually used to manage rabbit populations. Estimates of rabbit abundance were obtained monthly from roadside counts during a 25 month period developed at Donana National Park. Simultaneously, samples of the herbaceous layer were obtained from the area covered by censuses. Fluctuations in rabbit abundance were compared with changes in biomass and quality of the herbaceous plants. Rabbit abundance showed the best fit with the total protein availability in the herbaceous layer. Variations in the precipitation pattern between years, which affected both food availability and rabbit reproduction, and the incidence of the first epizootic of the rabbit hemorrhagic disease, caused the observed differences between maximum values of rabbit abundance in two consecutive years. The relative importance of other factors influencing rabbit numbers in Donana including myxomatosis, predation, and competition for food with large herbivores is discussed.
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We examined the influence of intrinsic (age, sex, and social status) and extrinsic (snow depth, snowpack hardness, temperature, available ungulate carcass biomass) factors in relation to time-activity budgets of coyotes (Canis latrans) in Yellowstone National Park, Wyoming. We observed 54 coyotes (49 residents from 5 packs, plus 5 transients) for 2507 h from January 1991 to June 1993. Snow depth, ungulate carcass biomass, and habitai type influenced the amount of time coyotes rested, travelled, hunted small mammals, and fed on carcasses. Coyotes decreased travelling and hunting and increased resting and feeding on carcasses as snow depth and available carcass biomass increased. Age and social status of the coyote influenced activity budgets. During times of deep snow and high carcass biomass, pups fed less on carcasses and hunted small mammals more than alpha and beta coyotes. Pups apparently were restricted by older pack members from feeding on a carcass. Thus, pups adopted a different foraging strategy by spending more time hunting small mammals. Coyotes spent most of their time hunting small mammals in mesic meadows and shrub-meadows, where prey densities were highest. Prey-detection rates and prey-capture rates explained 78 and 84%, respectively, of the variation in the amount of lime coyotes spent hunting small mammals in each habitat in each winter. Our findings strongly suggested that resource partitioning, as mediated by defense by older coyotes, occurred among coyote pack members in Yellowstone National Park.
Few concepts in ecology have been so influential as that of the trophic cascade. Since the 1980s, the term has been a central or major theme of more than 2000 scientific articles. Despite this importance and widespread usage, basic questions remain about what constitutes a trophic cascade. Inconsistent usage of language impedes scientific progress and the utility of scientific concepts in management and conservation. Herein, we offer a definition of trophic cascade that is designed to be both widely applicable yet explicit enough to exclude extraneous interactions. We discuss our proposed definition and its implications, and define important related terms, thereby providing a common language for scientists, policy makers, conservationists, and other stakeholders with an interest in trophic cascades.