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Ecol Res

DOI 10.1007/s11284-012-0949-1

O R I GI N A L A R T IC L E

Pedro Pereira Antonio Alves da Silva


Joana Alves Milena Matos Carlos Fonseca

Coexistence of carnivores in a heterogeneous landscape: habitat


selection and ecological niches

Received: 15 June 2011 / Accepted: 29 March 2012


 The Ecological Society of Japan 2012

Abstract Understanding distributional patterns and


mechanisms used by species for habitat selection is crucial
to adopt eective land management policies in terms of
biodiversity conservation. A heterogeneous landscape may
allow coexistence of species. That coexistence will be
dependent on the availability of the resources in the habitat
that has to be sucient to full their basic needs. The
present study aimed to investigate habitat selection, niche
breadth and niche overlap of three sympatric carnivore
species (Vulpes vulpes, Genetta genetta and Martes foina) in
a typically fragmented landscape from Central Portugal,
using camera-trapping techniques. The results obtained
revealed that the investigated species use the available
habitats dierently and in a non-random way. The red fox
showed the most specialized behaviour, positively selecting
coniferous forests. The common genet preferred eucalyptus, avoiding old-growth mixed woodland, in contrast with
stone marten that exhibited a strong preference for this
late habitat, avoiding eucalyptus. Concerning the niche
breadth, the genet had the highest value while the red fox
had the lowest one. The results obtained at the camera-trap
level showed that the highest niche overlap occurred between the genet and the stone marten which suggests that
these species can coexist and share the available resources.
Regarding the habitat level, the greatest niche overlap was
found for the stone marten and the red fox, indicating the
exploration of the same general habitat conditions by both
species. The results obtained in the present study support
the concept that landscape complexity allows coexistence
between species within the same trophic level.
P. Pereira J. Alves M. Matos C. Fonseca
CESAM and Department of Biology, University of Aveiro,
Campus de Santiago, 3810-193 Aveiro, Portugal
Present address:
A. Alves da Silva (&)
IMAR-CMA, Department of Life Sciences,
University of Coimbra, Largo Marques de Pombal,
3004-517 Coimbra, Portugal
E-mail: antonioalvesdasilva@gmail.com
Tel.: +351-239-855760
Fax: +351-239-855789

Keywords Habitat selection Camera-trapping


Sympatric carnivores Niche breadth Niche overlap

Introduction
Habitat destruction and fragmentation has been pointed out as the main threats to biodiversity (Wilcove
et al. 1998; Debinski and Holt 2000). For anthropogenic and natural reasons the availability of habitats
has been reduced and consequently, the proportion of
individuals of certain species in a particular habitat
type may be altered. Thus, population size and growth
rate may vary with the proportion of available habitats
of dierent types (Pulliam and Danielson 1991). Habitat selection can be described as the behaviour of a
given species, mediated by biotic and abiotic factors,
that determines environment eectiveness for that
species. In theory, the eectiveness of a habitat is
determined by the response of the organisms, the most
suitable habitat being that where survival and reproductive success are highest (Orians and Wittenberger
1991). Therefore, the study of habitat selection patterns
of wild species is crucial to understand which environmental variables inuence the tness of individuals
and the viability of populations (Wiens 1976; Fernandez et al. 2007; Benson and Chamberlain 2007; Pulliam
and Danielson 1991).
Habitat changes and destruction may be caused by
several processes that lead to the reduction of habitat
amount (habitat loss) and size of patches and also to the
increase of the number and isolation of patches (habitat
fragmentation) (Fahrig 2003). Furthermore, changes in
one habitat may easily modify properties of the surrounding habitats (van den Berg et al. 2001). Since the
habitats are often linked and dependent, habitat changes
may aect the entire ecosystem function.
In the Iberian Peninsula natural forests, originally
dominated by the oak species Quercus spp. (Ramil-Rego
et al. 1998), were converted into agricultural elds due to
human settlement, 7,0008,000 years ago (Bignal and

McCracken 2000). These alterations have continued,


and in the last century the remaining native forests were
gradually replaced by forests dominated by faster
growing and economically protable species, such as
maritime pine Pinus pinaster and eucalyptus Eucalyptus
globulus (Paiva 1998; Vieira et al. 2000). In Central
Portugal, the occupancy of pine and eucalyptus trees
represents actually 48.2 and 30.4 % of forest territory,
respectively (DGRF 2007).
The common genet Genetta genetta, the red fox
Vulpes vulpes and the stone marten Martes foina are
three abundant carnivores of the Iberian fauna, considered of low concern in terms of conservation
(Cabral et al. 2005). Despite their sympatry, several
studies have shown that they exhibit distinct patterns of
habitat use. The common genet has been described as
occurring predominantly in areas with considerable tree
and shrub densities (Palomares and Delibes 1994; Virgos et al. 2001; Galantinho and Mira 2009), while the
red fox presents a generalist/opportunistic behaviour in
habitat use. As so, red fox commonly use agricultural
elds like vineyards and olive-yards (Lucherini et al.
1995) or areas with dense forest cover, generally
selecting heterogeneous areas (Cavallini and Lovari
1994; Cagnacci et al. 2004). The stone marten has been
described as a generalist species (Virgos and Garc a
2002; Santos and Santos-Reis 2010), occurring in cork
woodlands, riparian vegetation, pasturelands, orchards
(Santos and Santos-Reis 2010) and in other cultivated
elds like olive-yards and vegetable-gardens (SantosReis et al. 2005) and scrubs (Rondinini and Boitani
2002).
Spatial complexity of the environment may foment
coexistence, habitat selection being one of the main
processes that promotes competitive coexistence
(Rosenzweig 1981). Dierent patches in landscape usually support a set of resources that allows sympatry of
species within the same trophic level (Levin 1974) and
enhances the persistence of predatorprey interaction
(Fahrig 2003). Interactions at the same trophic level are
favoured when species with dierent selective behaviour
coexist (Pimm and Rosenzweig 1981; Rosenzweig 1987).
In other words, cohabitation in a fragmented habitat is
privileged when specialist and generalist species are
present.
It is important to recognize and understand how
much the human alterations in landscapes may aect
wildlife occurrence and behaviour. Thus, the present
study aimed to expand the pre-existent knowledge
concerning the mechanisms for the selection of habitat
types and the coexistence of the sympatric red fox,
common genet and stone marten, focusing on the following aspects: (1) density and habitat selection among
the dierent landscape units present in a study area; (2)
niche breadth of each species and (3) niche overlap
between the three species. In addition, the adequacy of
current land-use policies in Portugal was discussed,
framed on the carnivore and overall biodiversity conservation.

Methods
Study area
The study area is located in Central Portugal, between
40 18 N40 27 N and 08 20 W08 25 W and
centred at the Bussaco national forest (altitude between
190 and 547 m a.s.l.). The climate of this region is predominantly Mediterranean, with some Atlantic inuences, characterize by hot summers (maximum 39 C)
and mild winters (minimum 1 C). Annual rainfall
is on average about 804 mm, mostly concentrated in
autumn and winter. The Bussaco forest (that represents
1 % of the study area) constitutes a microclimate, particularly fresh and rainy, with an annual precipitation of
about 1,525 mm.
Habitat classication
A GIS database was built for the study area to delineate
soil use classes, using ESRI software ArcGIS 9.3.
Based on aerial photos, eld surveys and COS 90 [landcover map produced by Instituto Geograco Portugues
(SNIG 1995)], six land-cover units were recognized in
the sampling areas: agricultural elds (composed by
traditional and more intensive agriculture, where the
main crops are corn Zea mays, bean Phaseolus spp.,
grape Vitis vinifera, olive Olea europea and potato
Solanum tuberosum); eucalyptus forest (Eucalyptus
globulus); coniferous forest (mainly Pinus pinaster but
also Pinus pinea); broadleaves forest (mostly oak species:
Quercus robur, Quercus rotundifolia and Quercus suber;
and other deciduous trees like Castanea sativa); urban
areas and Bussaco forest. The Bussaco national forest is
a wall fenced 105 ha old-growth mixed woodland, which
represents a relevant dendrological collections in Europe
(Paiva 2004). This forest started to be planted during the
seventeenth century, and some of the most predominant
species of trees and shrubs are Cupressus lusitanica,
Quercus robur, Q. pyrenaica, Q. suber, Castanea sativa,
Laurus nobilis, Arbutus unedo, Viburnum tinus, Ilex
aquifolium, Ruscus aculeatus, Phillyrea latifolia, Prunus
laurocerasus.The land-cover of the study area is quite
heterogeneous. Large extensions of monocultures of
eucalyptus (42 %) and pine (22 %), agricultural elds
(27 %), urban areas (5 %), shrublands (2 %), broadleaves forest (1 %) and Bussaco forest (1 %) feature
that mosaic.
Sampling areas
From the total area (16,470 ha) previously dened for
line-transect surveys of carnivore eld signs, three sampling sub-areas were selected (A, B and C) (Fig. 1),
totalizing 1,545 ha of eective sampling area. This area
was calculated using a buer of 600 m radius (approx. the

Fig. 1 Map of the study area in mainland Portugal, highlighting


the three sampling sub-areas (AC), the location of the camera
traps (black circles) and the main land-cover units. Sub-areas were

dened based on the predominant land-cover units: A is dominated


by eucalyptus, B by mixed forests and C by coniferous trees

mean distance between nearest cameras) around each trap


station, so the animals within the sampling area have an
equal probability of being caught in relation to their distance from the cameras. The three sampling sub-areas
were established according to land-cover characteristics,
the sub-area A being dominated by eucalyptus, sub-area B
by mixed forests (Bussaco forest included) and sub-area C
by coniferous trees (mainly maritime pine). In all of the
three sub-areas little and isolated patches of urban tissue
(around 4 % per sub-area) and agricultural elds occur
(around 10 % per sub-area).

played in all photos. The sensitivity level chosen was


normal or low depending on the sun and heat
exposure of each trapping-station. Each camera was
charged with a 2 GB SD card that allowed storing about
1,900 photos (with 8 MP resolution each), which corresponds to 633 shots approximately, and eight alkaline
batteries that, according to the manufacturer, provide
6 months of continuous work.
Cameras were placed at a height interval of
0.150.25 m and at a 2 m (approx.) distance from the
lure. Sand with domestic cat urine and canned sardines
were used as lure in the trap-stations. The lure was
replaced, and the SD cards were emptied every 3 days
(Kauman et al. 2007). Vegetation within the eld of
view of the cameras was removed to avoid false shots.
Trap-stations were distributed using a random grid
placed in each sub-area. Therefore, the habitats were
sampled following a stratied random design, according
to their availability (Table 1).
The space between cameras was a compromise
between maximizing the sampling area and trying to
detect all the individuals of the target species in the area.
For this, the mean radius of the home range (HR) of all
species were considered as reference values [HR stone
marten 260 ha, radius 455 m (Santos-Reis et al. 2005);

Camera-trapping surveys
Each sub-area was surveyed during 30 days (30 trapnights), between 15 June and 13 September 2010, the
total survey period being 90 days. This short trapping
interval increases the chances that the population will
stay stable and closed over the study period (Otis et al.
1978; Karanth and Nichols 1998; Trolle and Kery 2003;
Silver et al. 2004).
Ten Bushnell Trophy Cams were used, set to a time
interval of 1 s and to take three photos at each contact.
The date, time, temperature and lunar phase were dis-

Table 1 Number of cameras and photos of red fox, common genet


and stone marten in the dierent habitat types
Number of
cameras

Bussaco forest
Broadleaves forest
Coniferous forest
Eucalyptus forest
Agricultural elds
Urban areas

3
1
8
11
3
2

Number of photos
Red
fox

Common
genet

Stone
marten

15
7
114
10
3
2

3
2
21
51
2
0

76
6
157
70
13
1

HR common genet 330 ha, radius 512 m (Santos-Reis


et al. 2005); HR red fox 282 ha, radius 474 m (Cavallini
and Lovari 1994); mean global radius 480 m] and the
cameras were spaced at intervals of 500600 m. After
the site selection and the cameras positioning in the eld,
the mean distance (standard deviation) among nearest
cameras was: 601 63 m for sub-area A; 594 79 m
for sub-area B, 585 65 m for sub-area C and
594 69 m overall.
A pilot campaign of 15 days was rst conducted to
assess the eciency of the lures and the eectiveness of
the cameras in the study area, the time period needed to
empty SD cards and to replace the lure, the battery life
and other technical details of the cameras. The pilot
study was conducted in the three sub-areas using 10
cameras (sub-area A, 4 cameras; sub-areas B and C, 3
cameras each).
Individual identication of specimens
Individual common genets and red foxes were identied
considering pelage patterns, body marks (ear cuts, scars,
broken tails, etc.) and biometric parameters. Photos
with distorted perspectives or without sucient clarity
to allow understanding pelage patterns or any other
particular mark were not considered for individual recognition. Identication of specimens was not performed
for the stone marten due to the lack of singular characteristics of individuals.
Data analysis
Individual capture histories were constructed using a
standard X-matrix format (Otis et al. 1978) for the
identied red foxes and common genets using the photographs of the camera traps. Data on capture histories
were analysed using the CAPTURE software (Otis et al.
1978; Rexstad and Burnham 1991), developed to
implement closed-population capturerecapture models.
The test for closure included in CAPTURE software
(Otis et al. 1978) was used to determine the demographic
closure of the population under analysis. Eight fundamental capture-recapture models were evaluated for

closed populations that assumed dierent sources of


variation operating in the probability of capture to
estimate the population size of both species and the
corresponding standard error. The sources of variation
incorporated in these models were time (Mt, capture
probability varied between occasions and was constant
across individual), behaviour (Mb, capture probability
varied with behavioural responses diering for animals
captured or not captured previously), heterogeneity (Mh,
capture probability was dierent for each individual)
and pairwise combinations of these sources of variation
in capture probability (Mtb, Mth, Mbh and Mtbh).
Besides the previous seven models, we also tted the null
model (M0, capture probability was constant across
individual and time) to our data. The most appropriate
model was selected using the goodness-of-t test,
between model test (M0 vs. Mt, M0 vs. Mb and M0 vs.
Mh) and the model selection algorithm (model score
equal to one corresponds to the best t model) (Otis
et al. 1978; Rexstad and Burnham 1991) provided by
CAPTURE software. The selected model was then used
to estimate the abundance and the capture probabilities.
The selection ratios and Bonferroni-adjusted 95 %
condence intervals for each habitat type (Manly 2002)
was calculated to analyse the patterns of habitat selection exhibited by the red fox, the common genet and the
stone marten. Given the lack of consistent terminology
in studies about habitat selection, the terms preference
and avoidance were used to indicate that some
resources or habitats are signicantly more or less
selected than others comparatively with their availability.
The selection ratio (wij) was calculated as:
oij
wij
;
1
pij
where oij is the proportion of observation in the habitat i
and pij is the proportion of habitat i available for species
j. A selective use of habitat occurred when wij diered
statistically from 1 (P < 0.05). When wij values were
signicantly higher than 1 there was a positive selection
(preference), when wij values were signicantly lower
than 1 there was a negative selection (avoidance). The
standard error (SE) of the selection index was:
s
oij  1  oij
SE wij
;
2
uij  p2ij
where uij is the total number of observations. The
standardized selection ratios ( Bwij ) were used for direct
comparisons between selection ratios of the dierent
species and calculated as:
wij
Bwij PI
:
3
i1 wij
A log-likelihood Chi-square test ( v2L ) was used to test
the null hypothesis which assumed that carnivores randomly used all habitat types proportionally to its
availability.

Using the observed distribution of the species within


the set of habitat types, the niche breadth of each carnivore was calculated according to the Hurlbert method
(Hurlbert 1978):
0

Hj P
I

o2ij
i1 pij

taking into account the availability of the resources. H


0
was standardized to HA :
0

Hj  pimin
H Aj
;
1  pimin
0

ranging from 0 (strong preferences for some resources)


to 1 (no preferences for any resource) and where pimin is
the smallest proportion of all the habitats.
The overlap of the resource use among the three
carnivores was analysed at station and habitat scales.
Station and habitat scales were evaluated at camera-trap
and land-cover levels, respectively, to infer the overlap at
individual and population scales. The overlap was
measured through the Piankas index of niche overlap
(Pianka 1973):
PI
i1 oj  ok
Ojk q
;
6
PI
PI
2
2
o
o
i1 j
i1 k
that is a symmetric measure of overlap between species
j and k, ranging from 0 (no resources used in common)
and 1 (complete overlap of resource use).
The habitat selection, niche breadth and niche overlap were calculated using the R statistical language (R
2.12.1, R Development Core Team 2010).

bility due to time (M0 vs. Mt: v25 = 2.008, P = 0.848),


behaviour (M0 vs. Mb: v21 = 0.563, P = 0.453) or
heterogeneity (M0 vs. Mh: v21 = 3.148, P = 0.076),
and consequently, the null model was not rejected in
favour of these. Using the built-in model selection
algorithm of CAPTURE software, the null model (M0)
was the best tted model for the common genet capture
data (model score: M0 = 1.00, Mh = 0.97, Mb = 0.40,
Mbh = 0.63, Mt = 0.00, Mth = 0.49, Mtb = 0.37,
Mtbh = 0.71).
The null model was then used to estimate the population size of red fox and common genet and other
capture parameters like the capture probability.
Population estimate: size and density
Trapping eort resulted in a total of 553 animal
photos distributed by the six studied habitats (Table 1). The population density estimates were
1.17 0.08 and 1.49 0.16 individuals km2 for red
fox and common genet, respectively. Species presented
higher or lower density estimates depending on the
habitat type that dominated each sub-area. The subareas B and C were the regions with the highest
densities of both red fox and common genet. It was
impossible to estimate the population size of the stone
marten due to methodological constraints. However,
considering the linearity found between the number of
photographs and the number of animals, sub-areas B
and C were regions with high densities of stone marten (Table 2).
Habitat selection

Results
Closure test and model selection
The statistical test for population closure was consistent
with the assumption that the red fox population and the
common genet population were closed during the survey
period (red fox: z = 0.271, P = 0.607; common genet:
z = 1.016, P = 0.155).
The between model test for the red fox population did
not reject the null model M0 in favour of any of the
alternative models that incorporate the dierent sources
of variation into the probability of capture (M0 vs. Mt:
v25 = 9.015, P = 0.108; M0 vs. Mb: v21 = 1.448,
P = 0.229; M0 vs. Mh: v21 = 2.832, P = 0.092). The
discriminant function used in the model selection scored
the model M0 as the model that best ts red fox capture
data (model score: M0 = 1.00, Mh = 0.92, Mb = 0.35,
Mbh = 0.63, Mt = 0.00, Mth = 0.28, Mtb = 0.40,
Mtbh = 0.79).
In the case of common genet population, the between
model test did not indicate variation in capture proba-

The selection ratios obtained for each carnivore species


showed that the habitat types were not randomly
used (red fox: v2L5 = 170.47, P < 0.001; common
genet: v2L5 = 34.26, P < 0.001; stone marten: v2L5 =
163.56, P < 0.001). Indeed, the results expressed in
Table 3 illustrate a positive selection of coniferous forest
and a signicant avoidance of eucalyptus forest and
humanized areas (agricultural elds and urban areas) by
the red fox. The eucalyptus forest was the preferred
habitat to common genet, while Bussaco forest and the
humanized areas were avoided by this species. For the
stone marten, the high positive selection of Bussaco
forest and coniferous forest contrasts with avoidance
behaviour towards eucalyptus forest and humanized
areas. In general, the three carnivore species tended to
avoid the humanized areas (Table 3).
The direct comparison between the habitats selected
by the three studied species, using the standardized
selection ratios (Fig. 2), revealed higher selectivity by the
red fox, followed by the stone marten. All the species
showed dierent selection patterns, exhibiting dierent
values of Bw in all the sampled habitats (Fig. 2).

Table 2 Number of photos, population estimates and capture probabilities of red fox, common genet and stone marten in the three subareas of the study area
Red fox

Common genet

Number Population
of photos estimate
Sub-area A 4
Sub-area B 30
Sub-area C 117
Total
151

1
8
8
18

0.1
0.9
0.6
1.2

(1.02.0)
(8.012.0)
(8.010.1)
(17.021.2)

Stone marten

Capture
Number Population
probability of photos estimate

Capture
Number Population Capture
probability of photos estimate
probability

0.500
0.354
0.417
0.370

0.361
0.133
0.354
0.275

25
11
43
79

6
10
8
23

0.8
4.8
0.9
2.5

(6.09.1)
(6.045.4)
(8.012.0)
(20.030.3)

7
114
202
323

ne
ne
ne
ne

ne
ne
ne
ne

Sub-areas were dened based on the predominant land-cover units: sub-area A is dominated by eucalyptus, sub-area B by mixed forests
and sub-area C by coniferous trees. Population estimates are represented as abundance SE (95 % condence interval)
ne Not estimated due to methodological constrains

Table 3 Habitat selection ratios by the red fox, common genet and stone marten
Red fox

Bussaco forest
Broadleaves forest
Coniferous forest
Eucalyptus forest
Agricultural elds
Urban areas

Common genet

Stone marten

Selection ratios

Selection ratios

Selection ratios

0.927
1.298
2.642
0.169
0.185
0.185

0.748
0.534
<0.001
<0.001
<0.001
<0.001

0.354
0.709
0.930
1.643
0.236
0.000

0.001
0.556
0.689
<0.001
<0.001
<0.001

2.196
0.520
1.701
0.552
0.376
0.043

<0.001
0.534
<0.001
<0.001
<0.001
<0.001

0.227
0.479
0.123
0.052
0.106
0.130

0.201
0.495
0.174
0.137
0.165
0.000

0.220
0.210
0.097
0.058
0.102
0.043

Selection ratios are represented as w SE. If w is signicantly higher than 1 there was a positive selection, if signicantly lower than 1
there was a negative selection. Values with P < 0.05 are signicant and highlighted in bold font

(a)

(b)

(c)

Fig. 2 Habitat standardized selection ratios proles of red fox,


common genet and stone marten in a heterogeneous landscape
constituted by Bussaco forest (Buss), broadleaves forest (Broad),

coniferous forest (Conif), eucalyptus forest (Eucal), agricultural


elds (Agric) and urban areas (Urban)

Niche breadth and niche overlap

lap obtained at the station level indicated that an additional spatial segregation is occurring at a ner scale
than that happening at the habitat level.

Standardized niche breadth HA diers among the


studied species, with the highest similarity between the
0
0
common genet HA 0:731 and the stone marten HA
0:660, which presented a more generalist behav0
0
iour (higher HA comparing to the red fox HA
0:442.
In terms of niche overlap, the results indicated the
highest overlap between the common genet and the
stone marten at the station level, but at the habitat level
the maximum overlap occurred between the red fox and
the stone marten (Table 4). The lowest values for over-

Discussion
Understanding distributional patterns and habitat
selection mechanisms of carnivores, that are species
located at the highest levels of food chains, may be
essential to build eective land management policies,
where biodiversity conservation is a keystone.

Table 4 Piankas index of niche overlap among the three carnivores, calculated at two dierent spatial scales

Station scale
Habitat scale

Red fox
versus
common genet

Red fox
versus stone
marten

Common
genet versus
stone marten

0.116
0.464

0.288
0.909

0.413
0.684

The index is a symmetric measure of overlap between two species


and ranges between 0 (no resources used in common) and 1
(complete overlap of resources use)

An overview to the calculated carnivore densities


generally revealed low values. However, densities were
higher than those presented in other studies carried out
in the Iberian Peninsula (Palomares and Delibes 1994;
Sarmento et al. 2009, 2010). High densities of a certain
species in a particular habitat may suggest a great carrying capacity, thus, an improved quality, revealing high
suitability for that species. Since the studied carnivores
are game species in Portugal, the low densities found in
the present study may be related with the hunting
pressure to which these species are subject in rural areas.
The present results showed clearly that the three
carnivores used the available habitats dierently and in a
non-random fashion, showing preference and avoidance
trends. The preference of red fox for coniferous woodlands observed in this study was in agreement with that
reported by Cavallini and Lovari (1991). Moreover,
Goldyn et al. (2003) observed that red fox forages
mainly in habitats with a high availability of rodents,
like wood mouse Apodemus sylvaticus, that constitutes
its main preys (Serani and Lovari 1993; Carvalho and
Gomes 2004). The results of the present study followed
the same assumption because the coniferous forest
shelters a considerable density of wood mouse (number
of captures of wood mouse per 100 trap-nights: 3.9 in
Bussaco forest, not estimated in broadleaves forest, 2.0
in coniferous forest, 1.5 in eucalyptus forest, 0.8 in
agricultural elds, not estimated in urban areas; Matos
et al., unpublished data). In addition, the coniferous
forests represent a large and continuous extension,
which allows high occurrence of the red fox (Virgos et al.
2002). We believe that the non-selectivity of Bussaco
forest (the habitat with the highest abundance of wood
mouse) may have two main explanations: (1) the little
patch size of Bussaco forest (105 ha) which can disable
the settlement of a red fox population (Virgos et al.
2002) due to its need for large areas (Cavallini and
Lovari 1994; Lucherini and Lovari 1996); (2) the presence of very dense and cluttered arboreal and shrubby
vegetation in Bussaco forest, which condition predation
process by reducing preys accessibility.
The common genet showed positive selection for
eucalyptus and avoided Bussaco forest, which was a
quite unexpected result. However, it is known that the
genet has large exibility in terms of habitat choice
(Calzada 2002), since dense shrub cover, great availability of food (namely wood mouse) and low risk of

predation are provided (Palomares and Delibes 1988,


1994; Zabala et al. 2009; Virgos and Casanovas 1997;
Virgos et al. 2001, 2002). In this case, these ecological
requirements are all fullled since eucalyptus from the
study area present considerable understory cover (e.g.
Erica spp., Ulex spp., Rubus spp.), oering refuge and
prey abundance for genets, and low occurrence of red
fox (as shown in the Results) that is a potential
predator of genets. Bussaco forest also presents the
mentioned conditions, though (1) as mentioned above
for the red fox, the fragment size may constitute an
important constraint for the occurrence of carnivores
with a large HR, particularly in areas of about 100 ha
(Virgos et al. 2002); (2) Bussaco presents old arboreal
vegetation, giving a dense cover of large and elevated
branches. This arrangement most probably allows an
arboreal behaviour by the genets, which may reduce
their soil activity and condition their occurrence in the
cameras; (3) Bussaco is located at the highest altitude of
the study area. Several studies have demonstrated that
genets avoid high altitudes (Virgos and Casanovas 1997;
Calzada 2002), even at the HR scale (Zabala et al. 2009),
being more widely distributed in lower altitudinal
zones (Virgos et al. 2001). The low preference of genets for coniferous forests can be explained by the
high occurrence of red foxes, that is a potential predator
and competitor, and by the low availability of refuge
sites.
The stone marten was the only carnivore that positively selected Bussaco forest, simultaneously with
coniferous. The structural complexity of Bussaco forest,
with natural cavities, dense understory and diversity of
trees with well-developed foliage provides simultaneously wealth of resources to exploit and refuge for
martens. Stone martens are generalist feeders (Such and
Calabuig 2003; Santos and Santos-Reis 2010) able to
exploit the vertical and horizontal components of the
habitat, due to their arboreal behaviour (Padial et al.
2002). Therefore, Bussaco may constitute an important
habitat due to its great diversity of feeding items (i.e.
small mammals, fruit, arthropods, etc.). This becomes
particularly relevant during summer (Lanszki et al.
1999) when fruit constitutes the main item in the diet of
stone marten (Brangi 1995; Santos-Reis et al. 2005).
Among the forested habitats of the study area, Bussaco
forest detains the highest abundance and diversity of
available fruit which should inuence the selectivity of
stone marten towards this woodland. Most probably its
preference for coniferous forest is also related to food
availability that, in this case, was predominantly constituted by small mammals. Stone marten can minimize
the risk of predation by red fox in the coniferous forest
by using vertical component of the habitat. Arboreal
habits have been also reported for wood mouse
(Montgomery 1980; Rosalino et al. 2010). The avoidance of stone marten to eucalyptus plantations may be
related to the presence of other competitors, namely the
common genet, which therefore, can restrict the distribution of stone marten (Santos and Santos-Reis 2010).

The three sympatric species showed consistent


avoidance behaviour towards highly humanized landcover units, agricultural elds and urban areas with
human activities, pressure and perturbation. Furthermore, a great number of domestic dogs usually occur in
urban areas, and dog attacks to some medium-sized
carnivores have been reported (Palomares 1990; Palomares and Delibes 1994). All these factors can explain
the avoidance behaviours observed.
Besides habitat selection, niche breadth and overlap
constitutes crucial parameters to understand the ecosystem functioning. The niche breadth which indicated
the highest habitat specialization by the red fox may be
due to low population densities, fact which diminish the
need of the red fox to exploit unfavourable habitats and
makes the species more specialized in the suitable ones.
In agreement with the previous analysed results, the
common genet presents the highest niche breadth, using
a greater number of habitats.
In terms of niche overlap the results showed, as expected, a greater overlap at the population level (habitat
scale) than at the individual or HR level (station scale).
In terms of habitat scale, there is a coexistence of carnivore populations due to the large patch sizes presented
by the dominant habitats of the study area. On the other
hand, the results obtained for niche overlap at station
scale suggest the occurrence of a resource partitioning
between individuals of dierent species, which may
illustrate a more direct competition or distinct resource
requirements.
The low proportion of niche overlap between red fox
and common genet suggests that these species are direct
competitors that avoid themselves and do not share
common resources (niche segregation). Regarding red
fox and stone marten, the results showed populations of
both species coexisting and exploiting the same habitats,
but individuals presenting spatial segregation. This may
corroborate the explanation of the stone martens ability
to use the vertical component of environment. Stone
martens and common genets exhibit a considerable
niche overlap suggesting that they can coexist and exploit similar habitats. This should be due to the elusive
character of both species (Thompson 2004), reected by
their need of refuge (understory).
In conclusion, this study demonstrates that ecologically similar species may coexist within the same geographic region, being favoured by landscape complexity
and translated by habitat diversity. At a larger scale,
species share the same habitats, whereas locally they use
resources dierently, reducing overall competition. The
results of the present study and particularly the obtained
densities reinforce the established concept that unreserved territories also play an important role in biodiversity conservation, since natural reserves per se will not
eectively do it so (Fischer et al. 2006; Franklin 1993).
From a conservationist perspective, the habitat fragmentation and monoculture exploitations cannot be
assumed as presenting negative eects over the studied
carnivore community. Nevertheless, it must be high-

lighted that eucalyptus stands must maintain some


structural complexity, like a preserved understory layer,
in order to sustain or promote the studied species, specially the common genet. Moreover, the occurrence of
dierent carnivore populations is possible due to the
presence of large patches of dierent forest habitats, with
sucient area to allow species settlement. Furthermore,
the studied species are widespread and described as
generalists, and are able to adapt to various habitats.
Acknowledgments The authors would like to thank to T. Natal-daLuz for English corrections, and to the anonymous reviewers who
made helpful suggestions on an earlier draft of this manuscript.

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