Global Ecology and Biogeography, (Global Ecol. Biogeogr.

) (2007) 16, 281–289

Blackwell Publishing Ltd

META- Winter severity limits red fox populations

ANALYSIS
in Eurasia
Kamil A. Bartot* and Andrzej Zalewski†

Mammal Research Institute, Polish Academy of ABSTRACT

Sciences, 17-230 Bialowieza, Poland
*Correspondence: Kamil A. Bartot, Mammal
Research Institute, Polish Academy of Sciences,
17-230 BialowieΩa, Poland.
E-mail: kbarton@zbs.bialowieza.pl
†Present address: School of Biological
Sciences, University of Aberdeen, Tillydrone
Avenue, Aberdeen AB24 2TZ, UK.
Aim We investigated geographical variation in the density of the red fox in relation
to climatic variables, habitat productivity and seasonality to identify those factors
that were the best predictors of fox density.
Location Published data on red fox abundance were collated from 69 locations
over Europe and Asia.
Methods Using generalized linear models and the information-theoretic approach,
we analysed the contribution of climatic measures (winter and summer tempera-
ture, mean snow depth and duration), primary productivity and seasonality indices
[based on the fraction of photosynthetically active radiation (FPAR) index] to
account for variation in red fox density.
Results Red fox density in winter ranged from 0.001–2.8 individuals km−2; the
average density was 0.21 individuals km−2. Variation in red fox density was best
explained by the winter temperature and seasonality. Density decreased with declining
winter and summer temperatures, increasing degree of seasonality and increasing
duration of snow cover. There was no relationship with habitat productivity.
Main conclusions Our results indicate that winter climatic conditions and sea-
sonality, but not habitat productivity, may limit red fox density in Eurasia. One
explanation for the limitation of the red fox population may be the fox’s physiological
capability to cope with abiotic conditions. Concurrently, the severity of winter may
lead to reduced availability of the fox’s prey. That, together with a shorter reproduc-
tive period may result in lower reproductive output as well as lower survival of adults
and cubs.
Keywords
Environmental conditions, FPAR, population limitation, population density,
seasonality, Vulpes vulpes, winter severity.

ranges vary from being locally abundant to rare. Several major

INTRODUCTION
Mechanisms affecting species distribution and abundance are
important in understanding changes in plant and animal
communities, especially in the context of climate change (e.g.
Hughes, 2000; McCarty, 2001). Both the distribution and abun-
dance of species are often linked and may also exhibit a positive
relationship (Gaston et al., 2000). Interspecific abundance–range
size relationships suggest that locally rare species tend to have
narrow distributions, whereas locally abundant species tend to
be widespread (Gaston et al., 2000). However, this relationship is
usually not linear but rather triangular, as species with broader
mechanisms may account for this triangular tendency, for
instance the differences in variation of body size and metabolic
demands between species with smaller and wider ranges (Brown
& Maurer, 1987). Variation in the abundance of widespread species
within their geographical range, however, may affect the inter-
pretation of interspecific abundance–range size relationships.
The most general pattern explaining intraspecific geographical
variation in abundance is an ‘abundance–central distribution’,
which suggests that species abundance tends to be greater towards
the centre of a geographical range and lower at the periphery
(Brown, 1995). This assumes that central sites probably meet

© 2007 The Authors DOI: 10.1111/j.1466-8238.2007.00299.x

Journal compilation © 2007 Blackwell Publishing Ltd www.blackwellpublishing.com/geb 281
K. A. Barton and A. Zalewski
physiological and ecological requirements much better than
peripheral sites (Brown, 1984). Beyond the fact that this pattern
is widely accepted, little is known about the details of the form of
such relationships, although structural details may form an
important link between pattern and mechanism (Gaston &
Blackburn, 1999). Therefore, the question of what factors affect
abundance is still under debate (e.g. Currie & Fritz, 1993; Johnson,
1998; Melis et al., 2006; Zalewski & J^drzejewski, 2006).
Many factors may affect animal abundance and distribution,
including abiotic (such as temperature, day length or seasonality)
and biotic (predation or competition) factors. A commonly cited
factor is habitat productivity (McLoughlin et al., 2000). If a
population is limited by food resources, home range size may
decrease and abundance increase with increasing productivity of
the environment (Carbone & Gittleman, 2002; Kowalczyk et al.,
2003). The relative importance of productivity, however, may
vary between species and environments, with other factors
becoming more important, such as the severity of winters at
northern latitudes (Forsman & Mönkkönen, 2003). Climate may
limit the density of animals directly through their physiology or
indirectly by affecting the availability of resources. In addition,
fluctuations in habitat productivity (seasonality) can result in a
shorter reproductive period and may influence animal fitness,
such that fitness may decrease as seasonality increases (Boyce, 1979).
All these aspects are potentially interrelated, and distinguishing
their relative roles is key to understanding which factors limit
distribution and abundance.
An ideal species for studying the influence of both biotic and
abiotic factors on abundance is the red fox (Vulpes vulpes L.). It is
one of the most widespread vertebrate predators in the world. Its
range covers most parts of Europe, Asia and North America, and
it has also been introduced into Australia (Harris & Lloyd, 1991).
Environmental conditions vary markedly over their range, such
that foxes occur in areas of low productivity and very severe
climate. Foxes can attain high densities, and may exert an influ-
ence on prey populations (e.g. Kurki et al., 1997) and also act as
Figure 1 Distribution of collected data on the density of the red fox (dots).
282 Global Ecology and Biogeography, 16, 281–289, Journal compilation © 2007 Blackwell Publishing Ltd
a vector of important zoonotic diseases (Lindström et al., 1994;
Kauhala et al., 2006). Consequently, knowledge of the factors
affecting red fox density is important both from a theoretical and
an applied perspective. Most studies have analysed factors affect-
ing red fox populations in rural and urban areas or agricultural
landscapes (e.g. Harris & Rayner, 1986; Goszczytski, 1989), and
only a few concern populations inhabiting natural, relatively
unaltered habitats (Lindström, 1989). To our knowledge, there is
no study analysing the macroecological processes limiting the
size of fox populations.
The northern limit of the geographical distribution of the red
fox is determined by resource availability, but winter conditions
may also have some effect (Hersteinsson & Macdonald, 1992).
Therefore, we expect that habitat productivity should be an
important element affecting the abundance of this species. The
aim of this study was to identify the environmental factors correlated
with geographical variation in red fox density, which may explain
possible mechanisms limiting fox populations. Specifically, our
major objectives were to test: (1) whether there is a positive rela-
tionship between local habitat productivity during the breeding
season with winter fox abundance, (2) which climatic factors acting
in winter through survival, or in summer affecting reproduction,
play an important role in limiting fox populations, and (3)
whether seasonality of the environment affects the fox density
through variation in the length of the reproductive period. As the
distribution and abundance of species are linked, knowledge of
factors influencing abundance of the red fox can help to predict
changes in the distribution of this species with climatic changes.
MATERIALS AND METHODS
Data collection
We collated published data on red fox abundance from 69 locations
in Eurasia (Fig. 1). Because in rural regions culling by humans is
a likely factor determining fox abundance (Heydon et al., 2000),
© 2007 The Authors

to reduce this potential effect, we chose natural or semi-natural
habitats to include in our analyses (mostly national parks or
nature reserves). Red foxes are difficult to census accurately (e.g.
Cavallini, 1994), and studies in western Europe and North America
usually give indices of fox abundance (e.g. scat counts, spotlight
counts or number of foxes killed by hunters; Voight, 1999;
Heydon et al., 2000; Webbon et al., 2004). Snow tracking is the
most common method of censusing foxes in eastern Europe and
Asia, however, this would limit our study range to areas where
snow occurs (eastern and northern Europe and northern Asia;
there is a lack of snow-tracking data from North America).
Snow-tracking indices were collected from 55 locations. To
extend the range of the data, we also used density estimates given
by authors based on sand tracking, radio tracking or combined
methods involving den counting and visual observation (for
details see Appendix S1 in Supplementary Material).
Snow-tracking indices (tracks km−1 day−1) can give a relatively
good estimate of density using Priklonsky’s formula (Priklonsky,
1965; J^drzejewska & J^drzejewski, 1998). The tracking index
was multiplied by a constant 0.5π to obtain the total length of fox
trails laid on an average 1 km2 during 1 day, and divided by the
average daily distance covered (from Goszczytski, 1986). This
method assumes that the animal moves within a circular area
along a random path. The constant 0.5π is a mean track length
within a unit square, assuming that each of the sides of the square
is crossed only once.
We took mean values if the figures for a particular location were
available for more than one year (n = 21). Mean value was also cal-
culated in a few cases (n = 6) where there were only minimum and
maximum values over several years of snow-tracking index available.
Since density estimates calculated by authors (n = 14) were
likely to be more accurate then these calculated by us from snow-
tracking indices (biased through our choice of the daily move-
ment distance value for example), we also included the density
estimation method as a factor in our analyses.
Environmental variables
For every location, we obtained values of climatic variables and
vegetation productivity indices. Data on temperature and snow
cover were taken from data sets with a 0.5° resolution included
in the Global Ecosystems Database version 2.0 (downloaded
from the NOAA National Geophysical Data Center website:
http://www.ngdc.noaa.gov/). Satellite-derived data on snow cover
(monthly snow cover and snow depth; Chang et al., 1993) con-
sisted of monthly averages spanning a period from October 1978
to August 1987 (we used the last eight complete years). From
these, we calculated two variables: mean snow depth (in cm)
from November to February, and mean length of period with
snow cover (in months, i.e. the period when snow cover was
> 2 cm). Data on temperature consisted of monthly surface
temperature averages from 1931–60 (a relatively stable period;
Leemans & Cramer, 1992). From this data set, we calculated
mean winter temperature (average monthly temperature from
November to February) and mean summer temperature (average
monthly temperature from May to September).
© 2007 The Authors
Global Ecology and Biogeography, 16, 281– 289, Journal compilation © 2007 Blackwell Publishing Ltd
Winter severity limits red fox abundance
For an ecosystem productivity index, we used monthly 4 × 4
km MODIS FPAR (fraction of photosynthetically active radia-
tion) data sets covering January 2000 to June 2004 (downloaded
from the MODIS and MISR website at Boston University:
http://cliveg.bu.edu/modismisr/). The FPAR represents the percent-
age of the absorbed radiation available for photosynthesis
(Knyazikhin et al., 1998; Myneni et al., 2002) and it is used for
estimation of primary vegetation productivity (e.g. Zhao et al.,
2005). FPAR values were averaged within a 10-km buffer for each
location; this allowed us to account for the heterogeneity of the
area, since for many studies we did not have exact locations or
study area size (see Melis et al., 2006). The value of summer FPAR
was calculated as a monthly average from May to September.
As an index of seasonality, we calculated the yearly variance of
the FPAR index. We found variance to be a better estimator
of seasonality for our purposes than the standard deviation (cf.
Ferguson, 2002; Herfindal et al., 2005) with regard to the linearity
of a relationship with log-densities of the red fox. Taking into
account the strong linear relationship of the FPAR variance to
latitude (r2 = 0.71), to make the seasonality index more meaningful
we scaled its values so that it would be equal to approximately 0
at the equator and 1 at the latitude 90° N.
Statistical analyses
Initially, to address the problem of spatial autocorrelation, we
conducted simple and partial Mantel tests (Legendre & Fortin,
1989). With a simple Mantel test, we assessed the strength of
spatial autocorrelation in fox density. Partial Mantel tests served
to examine the relationships of fox density (log-transformed to
normalize variance) with each environmental variable, while
controlling for spatial separation (Legendre & Legendre, 1998). For
the distance calculations, locations were projected in equidistant
conic projection to minimize polar distortion.
As we found that our data were not spatially autocorrelated, in
subsequent analyses we did not account for spatial relationships.
To evaluate factors important in predicting fox density we used
generalized linear models and the information-theoretic
approach (Burnham & Anderson, 2002). Given the residual devi-
ance, the gamma error distribution with a logarithmic link was
utilized for models. The model selection procedure followed that
of Burnham and Anderson (2002). A candidate set of models
included those with the main effects alone, as well as several
combinations of environmental variables. We checked the en-
vironmental variables for collinearity by calculating all pairwise
Kendall’s τ correlation coefficients. Since winter measures (tem-
perature, snow depth and duration) were intercorrelated, no
more than one was included in each model. Additionally, the
‘method’ factor, consisting of two levels — ‘estimate from snow-
tracking index’ and ‘author’s custom estimate’ — was included in
all models. The models were ranked according to the corrected
Akaike information criterion (AICc).
We developed a confidence set of models, for which the
evidence ratios (relative likelihoods) versus the best model were
greater than a cut-off value of 1/8 (Burnham & Anderson, 2002).
We quantified the relative importance of each parameter in the
283
K. A. Barton and A. Zalewski

Figure 2 Relationship between red fox density and latitude and Figure 3 Relationship between red fox density and the most
longitude over Eurasia. Decrease in density between 50 and 100° E important explanatory variables, winter temperature (upper plot)
matches the Ural mountains region. Lines show the smoothing and seasonality index (yearly variance of FPAR index, lower plot).
splines fitted to the points (d.f. = 3 and 5, respectively). Lines show predictions by bivariate GLMs.

Table 1 Mantel r coefficients and P values (two-tailed) for the

confidence set of models by summing the Akaike weights over
all of the models in which the parameter of interest appears
(Burnham & Anderson, 2002).
All analyses were conducted using R 2.3.0 (R Development
Core Team, 2005).
results of simple and partial Mantel tests for the red fox density data.
Densities were log-transformed. The first row shows the results of a
simple Mantel test for spatial correlation of red fox density and the
remaining rows show results for partial tests of correlation of red fox
density with environmental variables while controlling for spatial
autocorrelation

RESULTS Mantel r P
−2
Red fox density averaged 0.21 individuals km (range 0.0001–2.8 Space 0.076 0.289
individuals km−2). An initial examination of the data indicated an Productivity 0.053 0.415
exponential increase in density with declining latitude (Fig. 2a) Seasonality 0.284 0.001
and a nonlinearity with longitude, with a clear decline between Snow depth 0.044 0.468
50° E and 100° E coincident with the position of the Ural Snow duration 0.237 0.003
mountains (Fig. 2b). Summer temperature 0.063 0.327
Red fox density was related to the examined climatic variables Winter temperature 0.260 0.001
and seasonality. Bivariate correlations showed that fox density
(log-transformed) increased with increasing winter (Pearson’s
r = 0.56, P < 0.001) and summer temperature (r = 0.43, P < summer productivity (r = −0.30, P = 0.01), also with mean yearly
0.001), and decreased with increasing snow duration (r = −0.57, productivity (r = 0.46, n.s.).
P < 0.001), snow depth(r = −0.41, P < 0.001) and seasonality (r = The Mantel correlation of log-transformed fox density with space
−0.58, P < 0.001) (Fig. 3). The weakest correlation was with was not significant (Table 1). The significance of the correlation

© 2007 The Authors

284 Global Ecology and Biogeography, 16, 281–289, Journal compilation © 2007 Blackwell Publishing Ltd
 Winter severity limits red fox abundance

Table 2 The confidence set of generalized linear models predicting

red fox density in Eurasia, ranked by corrected Akaike information
criterion (AICc). ∆i is the difference between the AICc of the best
model and that of model i and ωi is the Akaike weights. See Methods
for an explanation of the variables. The most parsimonious models
are at the top of the list. Model formulae shown do not include the
‘estimation method’ factor, which was present in all the models

Model Deviance AICc ∆i ωi

Seasonality + T.winter 104.57 −175.7 0.0 0.50

Seasonality + T.summer + T.winter 103.80 −173.9 1.8 0.21
Seasonality + T.winter + Productivity 104.57 −173.3 2.4 0.15
T.winter + Productivity + T.summer 106.23 −172.0 3.7 0.08
Seasonality + T.winter + 103.58 −171.6 4.1 0.07
Productivity + T.summer

T.winter, winter temperature; T.summer, summer temperature.

Figure 4 Predictions of the averaged model explaining

Table 3 Model averaged parameter estimates for the confidence set geographical variation in red fox density (Table 3), within actual
of five most parsimonious models predicting red fox density in ranges of the two most important explanatory variables, the winter
Eurasia, with unconditional standard errors (SE) and unconditional temperature and seasonality index. The remaining variables were
confidence intervals held constant at their mean values.

95% CI
DISCUSSION
Parameter Estimate SE Lower Upper
Limitations of methods
Intercept 2.589 1.334 −0.95 6.13
Winter temperature 0.088 0.170 0.03 0.15 When relating estimates of fox density to various satellite-derived
Seasonality − 5.721 1.683 −11.36 −0.09 indices we encountered several issues that were likely to influence
Summer temperature 0.025 0.215 −0.07 0.12 our results. FPAR has been used as a measure of net primary produc-
Productivity − 0.003 0.093 −0.02 0.01 tivity (Sellers et al., 1997; Ciais et al., 2005), and is closely related to
Method (snow tracking) − 0.817 0.652 −1.66 0.03 the normalized difference vegetation index (NDVI) (Veroustraete
et al., 1996; Myneni et al., 2002). Primary productivity, however,
may not accurately reflect food supply for foxes, as the availability
of rodents and hares (the main fox prey) may be limited by other
of each environmental variable with density did not change factors, like winter conditions and habitat structure.
when controlling for spatial relationship (partial Mantel test). The difference in time span over which the data on fox density
Three predictor variables were significantly correlated with density: were collected and the time at which the environmental variables
winter temperature, seasonality and snow duration. were measured may be important for the density data, as fox
The collinearity of the predictor variables was weak to modest numbers can fluctuate widely. We also had to assume that the daily
(Kendall’s τ: range 0.02 – 0.49), except between the winter variables distance moved by the fox does not differ between locations, as
(duration of snow cover, mean winter temperature and snow we did not have better figures. A fox’s daily movement distance
depth, τ = 0.54 –0.68). To reduce the effect of multicollinearity, may vary geographically, though, similarly as in the pine marten
we used only one of the winter variables in one model of the (Zalewski et al., 2004). So far, there is no single proven reliable
candidate set. method for monitoring changes in fox density (Sadlier et al.,
The selected confidence set comprised five models, which 2004). Track counting is the most common method, yet different
included all the examined variables except snow depth and duration sizes of census area could result in different estimates (Gaston et al.,
(Table 2). The model-averaged coefficients for summer tempera- 1999). However, we were not able to quantify the effect of size of the
ture and productivity were not significantly different from 0 census area because for many sites authors did not provide this
within 95% confidence intervals (Table 3). The relative import- information.
ances of parameters in the confidence set of models were: winter
temperature = 1.00, seasonality = 0.92, summer temperature =
Factors limiting fox populations
0.35, summer FPAR = 0.29, and for both summer variables =
0.35. Fox density predicted by the averaged model is shown in Over a large geographical area the density and distribution of
Fig. 4. many species are generally related to food abundance or habitat

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Global Ecology and Biogeography, 16, 281– 289, Journal compilation © 2007 Blackwell Publishing Ltd 285
K. A. Barton and A. Zalewski
productivity (e.g. McLoughlin et al., 2000; Carbone & Gittleman,
2002). However, in contrast to our expectation, red fox density
was not related to habitat productivity, which we used as a measure
of food availability, but rather to seasonality. Seasonality is
defined as the annual fluctuation of environmental conditions
and primary productivity. It is an important evolutionary pressure
affecting the life history and behaviour of mammals (Rosenz-
weig, 1968; Boyce, 1979; Lindstedt & Boyce, 1985; Andersson,
1994), and also affects reproduction because mammals living in
seasonal environments must give birth and raise young during a
short period of favourable conditions. There is a strong linear
relationship between seasonality and latitude, whereas such a
trend in primary productivity does not exist. We found that fox
abundance correlates with yearly variation in habitat productivity
much more strongly than with productivity itself. That, together
with the important role of winter temperature, suggests that the
winter food shortage or the variation in length of the reproduc-
tive period may be likely to limit the fox population. Seasonality
has been found to be negatively associated with age at maturity in
the moose (Ferguson, 2002); an analogous effect in the red fox
could limit the population through reproduction because young
vixens will not be able to reproduce in their second year.
Other observations also support the lack of a relationship
between density and productivity. For example, red fox home
range size, which is correlated with density (Kauhala, 2003), did
not relate to habitat productivity (Nilsen et al., 2005). Hersteinsson
and Macdonald (1992) analysed factors limiting the distribution
of red foxes at their northern limit and found that summer tem-
perature (taken as a habitat productivity index) predominantly
limited fox distribution. However, they also found that variables
related to winter conditions (minimum temperature, maximum
snow depth and duration of snow cover) had a significant effect
(Hersteinsson & Macdonald, 1992). Our study shows that fox
density declined exponentially with increasing winter severity
(especially mean winter temperature) and seasonality in habitat
productivity. Both of these studies point out the important role
that winter plays in limiting the abundance and distribution of
the red fox.
Winter may be a crucial period for a variety of reasons. Red
foxes are a medium-sized predator, active all winter (Harris &
Lloyd, 1991). The species-specific physiological tolerance to
abiotic conditions may be especially important in the north. The
physiological capability of individuals to cope with severe winter
conditions may determine abundance by increasing adult mor-
tality. A shorter reproductive period lowers the condition of the
cubs at the beginning of winter and also increases their mortality.
Winter mortality may be higher because food availability
decreases during harsh winters. The severity of the winter was
the most important factor affecting fox diet and foraging
(J^drzejewski & J^drzejewska, 1992). First, this may be because
rodents often stay under the snow and become less accessible to
foxes, with the percentage of voles in a fox’s diet decreasing in
deeper snow (Sonerud, 1986; Lindström, 1989; J^drzejewski &
J^drzejewska, 1992). Interestingly, however, we found that snow
depth had the weakest influence on fox density. Secondly, winter
harshness may affect the survival of voles as well as alternative
286 Global Ecology and Biogeography, 16, 281–289, Journal compilation © 2007 Blackwell Publishing Ltd
prey, reducing their winter abundance (Cederlund & Lindström,
1983; Lindström & Hornfeldt, 1994; Korslund & Steen, 2006),
however, it may increase the availability of ungulate carcasses
(Selva et al., 2005). On the other hand, the abundance of some
ungulates decreases with increasing winter severity (Melis et al.,
2006), which may reduce the availability of this type of food in
northern regions.
These results highlight the importance of winter (especially
winter temperature) as a possible bottleneck for red fox populations
by increasing mortality (due to physiological capability and/or
food shortage). Winter mortality may limit fox populations, and
indeed some studies have shown that high mortality in red foxes,
caused by, for example, intense culling by gamekeepers, can sup-
press fox populations (Heydon et al., 2000). However, winter
mortality may be compensated by higher reproduction, and a
few studies have suggested that compensatory density-dependent
reproduction may occur in the red fox (Cavallini & Santini, 1996,
Voight, 1999). In areas with higher mortality, the reproductive
output of vixens is much higher than in regions with lower
mortality (Voight, 1999). Therefore, summer red fox density may
not be related to winter conditions but to food abundance or social
regulation during the reproductive period. A full explanation of
the limitation of the fox population requires winter mortality as
well as summer reproductive compensation to be taken into
account.
Reproductive output is defined by the number of cubs per litter
and the number of reproductive females. In Europe, the average
number of placental scars in foxes varies between 3.2 to 7.3, but a
much greater variation is found in the percentage of barren
females (4–79%; Cavallini & Santini, 1996). After a harsh winter,
vixens are in a worse condition and therefore the number of fox
litters (reproductive females) is low (Kjellander & Nordström,
2003). Furthermore, the duration of winter and spring weather
determine the availability of food in spring and summer
(Pettorelli et al., 2005). Thus, winter harshness may have a direct
and indirect impact on reproductive output by reducing female
condition and food abundance during the reproductive period,
therefore limiting the fox population.
Red fox abundance decreased in higher latitudes, which is in
agreement with the abundance–central distribution hypothesis.
On the other hand, we found that densities of red fox increased
towards the south, which does not follow the predictions of the
abundance–central distribution hypothesis and previous findings
that population densities are generally smaller at lower latitudes
(e.g. Johnson, 1998). This result may be related to the fact that
our data covered only part of the southern range of red foxes.
Our findings have two important implications concerning
fox–human conflicts and competition with other medium-sized
predators. First, the red fox is a globally important species
because of its impact as a disease or parasite vector (Deplazes
et al., 2004). Most of these conflicts arise in modified habitats
whereas our analyses considered natural habitats. However, as
suggested by intraspecific abundance–occupancy relationships,
increasing the density of foxes in natural habitats with climatic
warming would also increase the range of occupancy of various
habitats. Furthermore, as dispersal between populations
© 2007 The Authors

increases, the transmission of disease or parasites may also
increase (including human-modified habitats). For example,
transmission of the fox tapeworm Ehinococcus multirocularis
which causes fatal liver infection in humans, is currently recognized
as major problem in many areas in Europe (Deplazes et al., 2004,
Graham et al., 2005). Secondly, further temperature rises can
influence a variety of ecological processes (Stenseth et al., 2002),
which would be most distinct at high latitudes in winter. In
northern Europe the red fox predates on pine martens, reducing
their density (Lindström et al., 1995). The red fox also limits the
southern range of the arctic fox by interspecific competition
(food and dens) (Hersteinsson & Macdonald, 1992; Elmhagen
et al., 2002; Tannerfeldt et al., 2002). Further climate warming
may promote an increase in abundance and expansion of the red
fox to new areas, and increase the negative influence on abun-
dance of the pine marten or such an endangered species as the
arctic fox.
ACKNOWLEDGEMENTS
The authors want to thank Xavier Lambin, Alex Douglas and two
anonymous referees for their valuable comments that improved
this paper, and Claudia Melis and Bernadetta Zawadzka for their
help with GIS. The work was supported by a Marie Curie EU
fellowship to A.Z.
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Global Ecology and Biogeography, 16, 281– 289, Journal compilation © 2007 Blackwell Publishing Ltd
Winter severity limits red fox abundance
BIOSKETCHES
Kamil BartoN is currently doing a PhD study on the
impact of vegetation productivity on rodent populations
in grasslands. His interests also include ecological
modelling and the use of the R language in statistical
computing.
Andrzej Zalewski has completed a PhD on the ecology
of pine martens in BialowieΩa National Park. He has also
conducted field studies on the use of space and
reproductive strategies in solitary carnivores (mainly
medium-sized mustelids). His current research is focused
on the influence of introduced American mink on native
fauna and its competition with native predators.
Editor: Kate E. Jones
SUPPLEMENTARY MATERIAL
The following supplementary material is available for this article:
Appendix S1 Data on estimates of population density of the red
fox (Vulpes vulpes) used in this paper.
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http://www.blackwell-synergy.com/doi/abs/10.1111/
j.1466-8238.2007.00299.x
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