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Piza Roca et al. / Lutra 2014 57 (2): 87-109 87
Introduction
e European badger (Meles meles) is present
in almost all European countries, from the
British Islands eastwards to the west bank of
the River Volga (gure 1). e species belongs
to the family of Mustelidae, in the order of
Carnivora. Recent studies showed that the
genus Meles includes several distinct species,
while this was considered only one species in
the past, the Eurasian badger. Accordingly,
the European badger is now described as a
distinct species from the Asian badger (Meles
leucurus) and the Japanese badger (Meles
anakuma) (Abramov 2001, 2003, Wozen-
cra 2005, Abramov & Puzachenko 2005,
2006). e Asian badger occurs from the
east of the Volga River to China and Korea,
till the border of the distribution of the Euro-
pean badger throughout the Lower and Mid-
dle Volga and the interuves of the Volga
and Kama (Abramov & Puzachenko 2006).
A clear geographic border in the northern
Caucasus between Meles leucurus and Meles
meles has not yet been clearly dened, as they
Environmental drivers of the distribution and
density of the European badger (Meles meles):
a review
Carme Piza Roca1,2, Maurice J.J. La Haye1,2 & Eelke Jongejans1
1Radboud University Nijmegen, Institute for Water and Wetland Research, Animal Ecology and
Ecophysiology group, P.O. Box 9100 (mail box 31), NL-6500 GL Nijmegen, the Netherlands,
email: carmepiza7@gmail.com
2Dutch Mammal Society, P.O. Box 6531, NL-6503 GA Nijmegen, the Netherlands
Abstract: Knowing the environmental factors aecting species’ distribution is an important basic step in ecologi-
cal research. Here, we present a literature review on the environmental factors inuencing European badger (Meles
meles) distribution and density. According to the published literature, the badger is a highly adaptive species, capa-
ble of using dierent environmental services and adapting to dierent conditions. However, when studying badger
populations across their distribution, a general pattern of preferred environmental characteristics and inuencing
factors arises. ese environmental characteristics may indicate the badger’s realised niche and may potentially
give an approximation of its fundamental niche. A combination of environmental factors favouring both sett (bur-
row) location and food availability appears to drive local badger success: terrain characteristics (both suitable soil
type for sett excavation and terrain heterogeneity for visual hiding), wood coverage and earthworm-rich grassland.
e presence and density of badgers vary depending on the geographic study area and the relative importance of
these specic environmental drivers. We discuss how these insights can assist spatial modelling studies, conserva-
tion and management, and future research on habitat suitability and badger density. We stress that more research
is needed to understand how each component of the life cycle of badgers is aected by environmental drivers, and
what the cumulative eect is on their spatial population dynamics.
Key words: environmental factors, European badger, fundamental niche, Meles meles, occurrence, review, sett.
© 2014 Zoogdiervereniging. Lutra articles also on the
internet: http://www.zoogdiervereniging.nl
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88 Piza Roca et al. / Lutra 2014 57 (2): 87-109
can occur sympatrically and may even repro-
duce, giving hybrids with mixed characters
(Abramov & Puzachenko 2007). e Japanese
badger occurs on the Japanese Islands (Bary-
shnikov et al. 2003). Finally, Del Cerro et al.
(2010) provide evidence for a fourth species of
badger named Meles canescens, distributed in
South-West Asia, from south of the Caspian
see and the Northern Caucasus to Tajikistan.
e taxonomic status of the badger nowadays
admits therefore four distinct species (Abra-
mov & Puzachenko 2013).
e European badger is a generalist, highly
adaptive, species which is capable of exploit-
ing a wide variety of habitats (Feore & Mont-
gomery 1999, Kauhala & Auttila 2010). It is
only absent from arctic zones, high altitude
regions and some islands (Griths & omas
1993). Analyses of the dynamics of an Eng-
lish population have shown that badgers start
breeding at an age of two years, that annual
juvenile survival (63%-77%) is lower than
adult survival (76%-88%), giving a genera-
tion time of 5.8 years, and that by an age of 7.3
years an average female has contributed half
of what she is going to contribute (through
reproduction) to population growth in her
life (Macdonald et al. 2009, van de Kerk et al.
2013). See gure 2 for a graphic representation
of the badger’s life cycle based on the study of
Macdonald et al. (2009).
e European badger is relatively abundant
in Europe, being only uncommon or present
in lower densities in the Netherlands, Bel-
gium, Estonia, Slovakia and Poland (Kranz et
al. 2008). Nevertheless, there is a general con-
cern about this species because it has showed
strong uctuations in numbers in many coun-
tries in the last century (Griths & omas
1993). In the 1970s and 1980s badgers obtained
Figure 1. Distribution range of the European badger (Meles meles), based on the distribution maps of Del Cerro et
al. (2010) and Abramov & Puzachenko (2013).
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Piza Roca et al. / Lutra 2014 57 (2): 87-109 89
a protected status in Britain, Ireland, Spain,
Portugal, Italy, Belgium, the Netherlands,
Albania, Greece, Estonia, Luxemburg and
Hungary (Griths 1991b). In 2008 the species
was ranked as Lower Risk/Least Concern on
the European Red List (Baillie & Groombridge
1996, Kranz et al. 2008), which means that pro-
tection had positive results. With its history as
a threatened species it is an interesting object
to formulate and study policy recommenda-
tions. erefore, knowledge is required about
the environmental factors driving the distribu-
tion and density of badger, which makes it pos-
sible to quantify habitat requirements, weigh-
ing management options, and assessing the
impact of habitat change. In broader ecological
studies, it is also of great importance to know
the interaction of the species of interest with its
environment in order to situate the species in
the ecosystem, its used niche and so on.
Figure 2. Matrix projection model for a European badger population near Oxford (Macdonald et al. 2009), based
on post-breeding census data. e rst stage therefore represents newborn cubs (zero-year olds). e population
model consists of ve parameters: age of rst reproduction (2), age of last reproduction (15), juvenile survival
(0.717), adult survival (0.837), and fertility (0.267).
Figure 3. Countries where the studies containing data relevant to the search topic were performed, i.e. the factors
inuencing distribution and density of the European badger as presented in this review.
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90 Piza Roca et al. / Lutra 2014 57 (2): 87-109
Table 1. Main environmental drivers of badger spatial population dynamics.
Environmental
driver
Components Contribution Potential eect on Eect Literature
Climate Rainfall range Higher seasonality Badger presence, spatial organisation, pop-
ulation size and density and group size.
Negative 1–3
Temperature range Higher seasonality Negative
Rain, temperature between 5– 15ºC
and high air humidity
Favours earthworm availability. Positive 4–8
Cold winters, dry summers and wind Decreases earthworm availability. Negative 9
Spring rainfall and high temperature Increase cub parasitic susceptibility. Negative
Terrain character-
istics and sett site
availability
Suitable geology Facilitates digging and drainage. Badger presence,
size and shape of occupied territory and
spatial organisation.
Limiting fac-
tor
10–21
Slope Favours earth removal.
Aspect Selection of sites on south to west-facing
slopes.
Terrain heterogeneity Visual hiding
Vegetation cover or small landscape
elements
Visual hiding
Groundwater level Sett building
Abandoned old setts Possibility for recolonisation Positive 22
Presence of habited setts Induces construction of new setts.
Habitat type Woodla nd Oers cover, shelter, structural support
for sett building and additional source of
food items.
Population density, group size and terri-
tory size.
Positive 1, 4–5,
10–15,
23–32
Hedgerows and scrub Cover and shelter
Grassland and pasture Foraging, source of earthworms.
Short grass is more suitable.
Arable elds: e.g. maize, wheat, barley Food source: cultivated food and earth-
worm content.
Food availability Large variety of items: e.g. earth-
worms, other invertebrates, birds’ eggs
and chicks, rodents, carrion, fruits,
cereals
Food source. Population size and density, territory size
and shape and spatial organisation.
Limiting fac-
tor
1, 11, 24,
29, 33, 43
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Piza Roca et al. / Lutra 2014 57 (2): 87-109 91
Built-up areas Urban infrastructure Decreases sett site availability.
Habitat destruction.
Fragmentation.
Badger presence, density and spatial
organisation.
Negative 2 6 –27,
44–48
Human population density. Disturbance on habitat. Population size and density. Negative
Agricultural land Habitat destruction.
Fragmentation.
Badger presence and spatial organisation.
Population size and density.
Negative
Urban and industrial areas Provide anthropogenic food sources. Population size and density. Positive 10, 14
Roads Increase mortality.
Habitat destruction. Fragmentation and
disturbance.
Badger presence and spatial organisation.
Population size and density.
Negative 14, 26,
45–55
Interspecic
competition
Mainly Vulpes vulpes, Nyctereutes
procyonoides but also other Carnivora.
Competition for sett sites and food. Distribution, population size and density. Potentially
negative, but
mostly neu-
tral
4, 2, 6,
56–71
Purposeful killingAllowed or regulated hunting and
poaching.
Increases mortality. Population size and density. In the most
severe cases also distribution.
Negative 5, 72–75
Diseases and para-
sites
Especially bovine tuberculosis (TB) Increases mortality and triggers poaching. Population size and density. In the most
severe cases also distribution.
Negative 76 –84
Other diseases (e.g. rabies) and para-
sites
Increases mortality. 85–88
Literature: 1. Virgós & Casanovas 1999a; 2. Kauhala 1995; 3. Johnson et al. 2002; 4. Kruuk 1989; 5. Kruuk 1978; 6. Griths & omas 1993; 7. Gerard 1967; 8. Bouché
1977; 9. Macdonald et al. 2010; 10. Huck et al. 2008; 11. van Apeldoorn et al. 2006; 12. da Silva et al. 1993; 13. ornton 1988; 14. Skinner et al. 1991a; 15. Matyástík &
Bicík 1999; 16. Doncaster & Woodroe 1993; 17. Neal 1972; 18. Neal 1986; 19. Broseth et al. 1997; 20. Macdonald et al. 1996; 21. van Moll 1999; 22. Roper 1992; 23. Feore
& Montgomery 1999; 24. Hofer 1988; 25. Palphramand et al. 2007; 26. van der Zee et al. 1992; 27. Schley et al. 2004; 28. Jepsen et al. 2005; 29. Kowalczyk et al. 2000; 30.
Palphramand et al. 2007; 31. Kruuk et al. 1979; 32. Kruuk & Parish 1977; 33. Kowalczyk et al. 2006; 34. Kruuk & Parish 1982; 35. Andersen 1954; 36. Neal & Cheeseman
1991; 37. Macdonald & Barrettn 1993; 38. Kruuk & Parish 1981; 39. Kruuk & Parish 1985; 40. Henry 1984; 41. Lüps et al. 1987; 42. Boyle & Whelan 1990; 43. Martín-
Franquelo & Delibes 1985; 44. Wright et al. 2000; 45. Aaris-Sørensen 1987; 46. Skinner et al. 1991b; 47. Mader 1984; 48. van Apeldoorn et al. 1998; 49. Bennett 1991; 50.
Clarke et al. 1998; 51. Vink et al. 2008; 52. Dekker & Bekker 2010; 53. Davies et al. 1987; 54. Harris et al. 1995; 55. Lankester et al. 1991; 56. Macdonald 1987; 57. Macdon-
ald et al. 2004; 58. Neal and Cheeseman 1996; 59. Fedriani 1993; 60. Kowalczyk et al. 2008; 61 Drygala et al. 2010; 62. Oerlemans & Koene 2008; 63. Goszczynski 1999; 64.
Kauhala & Auttila 2010; 65. Kauhala & Holmala 2006; 66. Jedrzejewska & Jedrzejewski 1998; 67. Martín et al. 1995: 68. Palomares & Delibes 1997; 69. Macdonald 2009;
70. Delibes & Calderón 1979; 71. Fedriani et al. 1999; 72. Grigorov 1987; 73. Cresswell et al. 1990; 74. Bego 1992; 75. Griths 1991a; 76. Gormley & Costello 2003; 77.
Olea-Popelka et al. 2003; 78. Krebs et al. 1997; 79. Gallagher & Clion-Hadley 2000; 80. Grin et al. 2005; 81. Donnelly et al. 2003; 82. Woodroe et al. 2006; 83. Dolan
1993; 84. WHO 1978-99; 85. Artois et al. 1991; 86. Schwierz & Wachendörfer 1981; 87. Harris & Yalden 2008; 88. Anwar et al. 2000.
Lutra_57_2_Text_v3.indd 91 06/02/2015 21:27
92 Piza Roca et al. / Lutra 2014 57 (2): 87-109
Methods
For this review we searched the ecologi-
cal literature for information about the fac-
tors aecting badger distribution and density
all around Europe, thus covering all dier-
ent environments that this animal inhabits.
We used the Web of Science and the search
engine Google Scholar. e search terms
were combinations of “European badger”,
“Meles meles”, and “habitat”, “preferences”,
“selection”, “environmental factors”, “driv-
ers”, “distribution”, “occurrence”, “density”,
“aect population”, “niche”, “sett”. Most of
the search was focused on papers published
in English and the search terms were always
in English. However, some data were found in
publications in other languages such as Span-
ish, French and Dutch. e time range of the
collected ndings is from 1970 until present.
We use this time frame rst because badger
populations across Europe had started to uc-
tuate from around the early 1970s, mainly due
to direct or indirect human pressure (Grif-
ths & omas 1993), oen becoming endan-
gered, and of global concern and thus ecologi-
cal studies on this species started to increase.
erefore, more data is available from then
on. Moreover, due to the rapid landscape
change all around Europe, studies performed
before these dates may not be applicable now-
adays. All factors that were reported to inu-
ence the spatial population dynamics of the
badger, from habitat composition to resources
availability and abiotic and biotic interac-
tions, are subsequently presented and dis-
cussed. We organised the driving factors in
several categories. First we present the abi-
otic factors in the landscape: climate factors
(such as temperature and meteorology), ter-
rain characteristics and factors determining
badger sett (burrow) site availability (such as
soil characteristics, slope and orientation and
groundwater level), habitat composition (such
as land use factors and landscape elements),
and nally food and food availability. Subse-
quently we analysed anthropogenic factors:
built-up areas and human density, roads and
Figure 4. European badger sett site selection for either woodland or other landscape elements. Each pair of col-
umns shows the results, respectively, from ecological studies in Essex (United Kingdom, Skinner et al. 1991a),
Luxemburg (Schley et al. 2004), Northern Moravia (Czech Republic, Matyáštík & Bićík 1999), Sudety Mountains
(Poland, Bartmańska & Nadolska 2003), and central Spain (Virgós & Casanovas 1999a). e rst column shows
the percentage of woodland present in the studied landscape (L). e second column shows the percentage of setts
(S) located in woodland.
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Piza Roca et al. / Lutra 2014 57 (2): 87-109 93
hunting. Finally we present the biotic factors
such as interspecic competition, diseases
and parasites.
Results
We found 96 studies from 18 dierent Euro-
pean countries providing relevant informa-
tion on the research topic. e UK had the
most performed studies (50 containing data
presented in this review), followed by Poland
(8), Spain (8), the Netherlands (7), Finland (6)
and Switzerland (6) (gure 3).
In the following sections we present an
overview of the most important factors we
found determining the distribution and den-
sity of badgers (see also table 1).
Climate
Climate is the best explanatory factor of
badger occurrence in dierent countries, e.g.
in Spain (
Virgós & Casanovas 1999a) and in
Finland (Kauhala 1995). Across Europe, John-
son et al. (2002) demonstrated that badger
group size decreases with rainfall range and
that badger density is negatively correlated with
temperature range (dierence between maxi-
mum and minimum temperature).
Hence, badgers are more abundant in rainy
regions, as rain favours the presence of earth-
worms in the soil (Kruuk 1989, Griths &
omas 1993). Climatic conditions are known
to be important for earthworm availability
especially during the night, when worms come
to the surface and badgers can forage on them
(Gerard 1967, Bouché 1977). Temperatures
between 5-15 ºC and high air humidity have a
positive inuence, while cold winters, dry sum-
mers and wind negatively inuence the presence
of earthworms (Kruuk 1978). us, climatic fac-
tors can aect badger populations indirectly by
inuencing earthworm availability.
In the United Kingdom, Macdonald et al.
(2010) showed that seasonality, through a
variation of temperature and rainfall, has a
complex inuence on badger populations.
Late-summer low temperatures and rainfall
have a signicant positive inuence on badger
survival, as cool and moist conditions favour
earthworm availability on the soil surface. On
the other hand, spring rainfall and tempera-
ture negatively inuence badger populations
as wet and warm conditions lead to higher
parasite susceptibility of the cubs. Finally,
colder winters generally result in badgers
staying underground and this reduced activ-
ity has a positive eect on survival due to
fewer badgers being hit by cars.
Terrain characteristics and availability
of potential sett sites
Location suitability for sett building may
determine the size and shape of badger occu-
pied territory (Doncaster & Woodroe 1993)
and spatial organisation within a region (da
Silva et al. 1993). A good sett site requires a
suitable soil to facilitate digging and drain-
age, such as sandy soils, in combination with
some gradient and vegetation cover (Neal
1972, Neal 1986, ornton 1988, Skinner et
al. 1991a, Good et al. 2001, Fischer & Weber
2003). Not only the presence of gradient but
also orientation of the slope is an important
factor inuencing sett location. is was an
important driver of sett site selection in Essex,
Norway and Northern Moravia: south to west
facing slopes were preferred most (Skinner
et al. 1991a, Broseth et al. 1997, Matyáštík &
Bićík 1999). On the other hand, it has also
been shown that badgers prefer terrain het-
erogeneity independently of slope orienta-
tion (ornton 1988, Macdonald et al. 1996).
Groundwater level can also be relevant for
sett excavation, as it is impossible to dig a sett
when the groundwater level it is too high (van
Moll 1999). erefore, in grounds with high
groundwater levels, this will be a constraint.
Hence, the location of badger setts is
selected according to the presence of favour-
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94 Piza Roca et al. / Lutra 2014 57 (2): 87-109
able conditions, which results in an hetero-
geneous distribution of setts in an area (van
Apeldoorn et al. 2006). Abandoned old setts
can be suitable for badger recolonisation, and
the presence of inhabited badger setts also
positively inuences the construction of new
setts (Roper 1992).
Habitat composition
Besides terrain topography, the type of habitat
(e.g. woodland, shrubs, pastures) also aects
badgers. Feore & Montgomery (1999) showed
a preference of badgers for sett sites at or near
the interface of two habitat types, especially
woodland or shrubs with pasture. Huck et al.
(2008) found that habitat type was the most
important factor explaining sett location in
urban areas: the presence of wood and scrub
mattered more than soil conditions and food
availability.
In the Netherlands, Apeldoorn et al. (2006)
found in a local study on habitat use and com-
position (between the towns of Utrecht and
Hilversum) that pastures were the main driv-
ers of badger distribution due to the provision
of food, followed by broadleaf forest which
was preferred for digging setts, mixed forest
and maize elds. ey also found badger setts
specically located near the edge of the for-
est, close to the grasslands and arable elds.
e presence of water was also important for
badger inhabitation. is suggests that this
habitat mosaic was selected to enhance both
sett building and food searching.
In England, the reproduction and weight
of badgers were higher in deciduous wood-
land than in any habitat type, such as pasture
(da Silva et al. 1993). is was a paradoxical
nding as pasture contains a higher biomass
of earthworms (Kruuk 1978, Hofer 1988),
which are one of British badgers’ main food
item. e higher weight of badgers in wood-
lands may have been the result of providing
an additional source of food items (e.g. car-
rion, blackberries and acorns), humidity
and protection from the wind (da Silva et al.
1993). Also in the United Kingdom, conifer-
ous woodland appeared to be more impor-
tant than previously thought. e positive
eect of woodland on the badger population
is likely due to the fact that woodland consti-
tutes a refuge from human activity and pro-
vides structural support for the construction
of setts within the root system (Palphramand
et al. 2007).
Woodland is in fact strongly preferred
by badgers for sett location and therefore
strongly inuencing badger density (gure
4). In England, wood density in the landscape
was positively correlated with sett density
(ornton 1988), which is taken as represent-
ative of badger density in most studies. is
eect was especially important in more open
landscapes in the Netherlands (van der Zee
et al. 1992). In Essex (United Kingdom), even
when 73% of the country is covered by arable
and pasture land, 87% of all setts were located
in woodland, hedgerows and scrub (Skinner
et al. 1991a). In Luxemburg 88% of setts were
located in forest while only 34% of the land
is covered by forest (Schley et al. 2004). e
most preferred forests were conifer and decid-
uous forest (38% of total setts each), followed
by mixed forest (12%). e other habitats with
setts were shrub (5%), hedgerow (3%), grass-
land (2%) and arable soil (0.3%).
In Northern Moravia (Czech Republic),
woodland was also the most preferred habi-
tat for sett construction, as 75% of setts were
located in this habitat type (Matyáštík &
Bićík 1999). e most frequented habitat was
mixed forest (33%), followed by coniferous
and deciduous forests (26% and 16% respec-
tively). Other setts were located in habitats
with rocks (11%) and only 6% in elds. In the
Polish Sudety Mountains badgers show a very
strong preference for woodland when build-
ing their setts, as 98% of setts were found in
woodland, and only 2% in the open areas,
although forests cover only 29% of the moun-
tains (Bartmańska & Nadolska 2003). Of the
setts located in woodland, 57% were found
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Piza Roca et al. / Lutra 2014 57 (2): 87-109 95
in deciduous and mixed forests, and 43% in
coniferous forests. In a study in central Spain
there was also a clear preference for wooded
places, as 50% of setts were located at sites
with >50% tree cover (Virgós & Casanovas
1999b). In Denmark forest cover, together
with terrain heterogeneity, were the most
important explanatory variables for sett pres-
ence using species distribution modelling
(Jepsen et al. 2005). Variation in wood cover
explained 22% of the total variation in badger
sett densities in Białowieża Primeval Forest in
Poland (Kowalczyk et al. 2000).
Small landscape elements like hedgerows,
orchards and small patches of woodland oer
coverage and favour badger sett location,
especially in agricultural land (Neal 1972,
ornton 1988). ese small landscape ele-
ments, however, have been removed to create
larger elds for agriculture in many regions,
which might negatively inuence badger pop-
ulations (ornton 1988).
Food and food availability
Badger territory size and density of setts in the
landscape are mainly shaped by food abun-
dance and availability (Kruuk 1989, Kowalczyk
et al. 2000). In areas with low or dispersed food
sources badgers move longer distances, cover
larger daily ranges and defend larger territo-
ries (Kowalczyk et al. 2006). Anyhow, badgers
behave as contractors, which means that they
keep a minimum territory where they can nd
just the sucient resources (Kruuk & Mac-
donald 1985). e mean size of group territo-
ries strongly diers between European regions,
from 0.14 km2 in the open habitats of the Brit-
ish Isles (Cheeseman et al. 1981) to 25 km2 in
the continuous woodlands of Poland (Kow-
alczyk et al. 2003). e amount of food avail-
able in the territory also strongly inuences
the number of individuals and group size in
badger populations (Kruuk & Parish 1982).
erefore it is important to know the common
food sources of this species.
e European badger’s diet is very variable.
is species is the most omnivorous mustelid,
an opportunistic forager that takes a large
variety of animal and plant food sources,
such as earthworms and other invertebrates,
birds’ eggs and young, rodents, carrion like
road kills, fruits, bulbs, acorns, oats and
wheat (Andersen 1954, Neal & Cheeseman
1991, Macdonald & Barrett 1993). In temper-
ate areas in Europe, like the British Islands,
earthworms are the main food source for
badgers (Kruuk & Parish 1981, 1985, Henry
1984, Lüps et al. 1987, Boyle & Whelan 1990).
But in drier regions, such as Spain, earth-
worms are not always available and badgers
specialise more in lagomorphs, mainly rabbits
(Oryctolagus cuniculus), and some fruits such
as olives and also arthropods (Barea-Azcón et
al. 2001).
In temperate regions, the main food items
for badgers consist of earthworms, espe-
cially Lumbricus terrestris, the whole year
and maize from arable elds during autumn
and winter (van Apeldoorn et al. 2006). As a
main food source, the abundance of earth-
worms can strongly inuence badger popula-
tions. Indirectly, the presence of earthworms
and the distribution of earthworm patches
have also been shown to aect the number of
badgers in a social group, the spatial organ-
ization (Kruuk & Parish 1982) and the con-
guration of badger territories (Hofer 1988).
Earthworms are common in pasture and old
forest. Acidity of grassy peatlands and for-
ested sandy soils lead to low earthworm bio-
mass densities, which results in poor condi-
tions for badgers. In Oxfordshire (England),
grasslands and broadleaf forests were proven
to oer relatively good food conditions, while
mixed and coniferous forest oered worse
food conditions (Kruuk 1978, Hofer 1988, da
Silva et al. 1993).
In England, barley, wheat and acorns were
shown to be secondary food items. Other food
sources eaten by badgers were insects, pig-
nuts, small mammals, birds, amphibians, car-
rion, etc. (Kruuk & Parish 1982). When feed-
Lutra_57_2_Text_v3.indd 95 06/02/2015 21:27
96 Piza Roca et al. / Lutra 2014 57 (2): 87-109
ing from cereals, badgers preferred wheat and
oats to barley (Kruuk 1989).
In the United Kingdom, territory size was
found to be negatively correlated with grass-
land proportion (Palphramand et al. 2007),
but it was positively correlated with the num-
ber of grassland patches. is suggests that
grassland is a key resource for badgers, likely
because it constitutes a source of earthworms.
However, grassland inuence depends on the
length of grass. Long grass was shown to be
unsuitable for badgers (Kruuk et al. 1979).
Badgers visit grasslands especially to forage
for earthworms (Kruuk & Parish 1977), and
catching them is much easier in short grass. A
thick soil cover, such as dead litter and vegeta-
tion, might also be more dicult to forage on
earthworms.
Type and amount of crops and grasslands
might therefore be important for badger pop-
ulations, as they directly provide various food
sources. Changes in agricultural land strongly
aect earthworm biomass (Edwards & Loy
1977, Kruuk 1978, Eijsackers 1983, Lofs-Hol-
min 1983) and can hamper earthworm avail-
ability, having a negative impact on badger
populations.
Built-up areas and human population
density
Many studies have found a negative correla-
tion between human population density and
badger sett density (e.g. Schley et al. 2004) and
also between urbanised area and sett density
(Wright et al. 2000). Urban areas, roads and
agriculture have been responsible for badger
population decline and distribution contrac-
tion throughout most of their geographic
range (e.g. Aaris-Sørensen 1987, Skinner et
al. 1991b). ese factors lead to habitat frag-
mentation, reducing the suitable habitat to
small unfavourable isolated patches (Mader
1984, van Apeldoorn et al. 1998), which can
no longer support sustainable badger (sub)
populations.
However, the anthropogenic transforma-
tion of the landscape may not always have a
negative impact on badger populations. As
demonstrated in Switzerland (Do Linh San
et al. 2011), an increase of agricultural land-
scape provides an additional food source and
badgers prot from this human-made food
rich habitat by adopting cereals and maize as
a main food item.
In the United Kingdom, Huck et al. (2008)
showed that badgers are capable of estab-
lishing relatively dense populations in urban
environments. ese provide some advan-
tages in providing anthropogenic food
sources. In Essex 15.9% of setts were indeed
found in urban and industrial areas, likely
due to badgers avoiding agricultural land and
searching for human-generated food (Skinner
et al. 1991a).
Roads
Roads may aect the distribution and popu-
lation size of badgers in three dierent ways:
1. ey constitute a barrier for badger move-
ment and dispersion causing habitat frag-
mentation. 2. ey increase badger mortal-
ity through trac kills, and 3. ey decrease
badger colonisation by producing disturbance
by higher human activity and trac noise
(Bennett 1991, Clarke et al. 1998).
e increase in number of roads and their
use was the major factor causing the his-
toric decline in the badger population in the
Netherlands (van der Zee et al. 1992) and is
the main cause of badger mortality nowa-
days in this country (Vink et al. 2008). In the
Netherlands, every year 10–20% of the total
badger population is killed on roads, mainly
in March and less in the winter months. Per
km of road most mortalities occur on pro-
vincial roads (Dekker & Bekker 2010). Miti-
gation measures have been shown to reduce
mortality of badgers (Vink et al. 2008, Dekker
& Bekker 2010). ese include construction of
passages and fences, reducing speed limits
Lutra_57_2_Text_v3.indd 96 06/02/2015 21:27
Piza Roca et al. / Lutra 2014 57 (2): 87-109 97
and closing critical roads.
In England, trac is also the major reason
for badger mortality, causing 49% of all mor-
talities (Davies et al. 1987, Harris et al. 1995).
In Surrey and Gloucestershire the impact of
road kills was even more dramatic: 59% and
66% respectively (Clarke et al. 1998). Road
kills in England show a strong seasonal var-
iation, with peaks in mortality occurring in
spring and late summer (Davies et al. 1987).
e relationship between road mortality and
trac load was found to be parabolic (Clarke
et al. 1998), possibly because badgers are dis-
couraged to cross the busiest roads (van der
Zee et al. 1992). erefore, major roads may
have mixed eects: a higher impact on badger
movements increasing the fragmentation
eect of roads (Lankester et al. 1991), but a
lower number of road kills as fewer badgers
attempt to cross such busy roads.
In Essex, sett density was signicantly
inuenced by road type and distance to roads.
e busiest roads were clearly avoided: signi-
cantly fewer setts than expected were found
within 10 m of a road than at 100–999 m from
a road (Skinner et al. 1991a). e size of the
badger population was also negatively corre-
lated with road and trac densities (Skinner
et al. 1991b).
Hunting
Hunting of badgers was a real threat for
badger populations all around Europe before
protection policies were established. Hunting
of badgers was a cause of population decline
for instance in Albania (Bego 1992), Bulgaria
(Grigorov 1987) and the United Kingdom
(Cresswell et al. 1990). Nowadays, hunting
of badgers is either strictly regulated or for-
bidden in the European countries where this
species is protected: the United Kingdom, Ire-
land, the Netherlands, Denmark, Belgium,
Italy, Greece, Spain, Portugal, Luxemburg,
Hungary, Estonia and Albania (Griths &
omas 1997). Nonetheless, in other Euro-
pean countries such as Poland (Mysłajek et
al. 2012), this animal is still seen as a small-
game hunting target or as a pest. is shows
that badgers are perceived very dierently
within Europe (Griths 1991a). In most of
the countries where hunting is allowed, this
is prohibited during the reproductive season
(Griths & omas 1993), but some coun-
tries oer very poor protection from hunting,
such as Finland and Austria, or no protection
at all, such as Bulgaria and Macedonia (Grif-
ths 1991a). In France and Germany hunt-
ing is popular but appears not to be a major
threat (Keuling et al. 2011, FDC 2014). Com-
pared to other mammalian game species, only
in Sweden, Switzerland and Norway the num-
bers of badgers legally hunted surpassed 4%
of the most popular mammalian game species
(Griths & Krystufek 1993).
Nonetheless, poaching seems to be a threat
to badger populations all around Europe
(Griths & omas 1993), especially in the
United Kingdom and Ireland (Cresswell et
al. 1989, Smal 1995). Illegal hunting prevents
the badger population in Albania to recover
(Bego 1992). In summary, hunting may
endanger badger populations in countries
where it is still allowed or seasonally allowed
or where it is practised illegally. For this rea-
son the Council of Europe (1987) has asked
all countries that allow hunting of badgers to
take measures to protect their stocks.
Interspecific competition
Red fox (Vulpes vulpes) is a potential competi-
tor of badger for food and sett sites because
it occupies a similar ecological niche (Mac-
donald 1987, Kruuk 1989). Nevertheless, both
species can apparently cohabit the same area.
Aggressive as well as peaceful encounters have
been reported (Neal & Cheeseman 1996), but
most encounters are not signicantly violent
and badgers take the dominant role (Mac-
donald et al. 2004). e two species have even
been found sharing the same sett (Macdonald
Lutra_57_2_Text_v3.indd 97 06/02/2015 21:27
98 Piza Roca et al. / Lutra 2014 57 (2): 87-109
1987, Fedriani 1993).
e raccoon dog (Nyctereutes procyonoides)
is an invasive species from Russia which has
already successfully colonised many north-
eastern European countries, including Fin-
land, Norway, Germany and Poland (Kau-
hala 1995, Kowalczyk et al. 2008, Drygala et al.
2010) and the species is likely going to inhabit
and quickly increase in other countries such as
the Netherlands in the near future (Oerlemans
& Koene 2008, Mulder 2012, Mulder 2013).
is species is also a potential competitor for
badgers for food and sett sites. Both species
are omnivorous and though raccoon dogs do
not construct burrows themselves, they oen
inhabit badger setts for reproduction and win-
tering (Goszczynski 1999).
Still, despite this ecological overlap, the rapid
invasion and growth of the raccoon dog popu-
lation in Finland has not been found to have a
negative eect on the native badger population
(Kauhala 1995). ey have been sympatric for
more than 50 years and both species increased
in number during this period (Kauhala & Aut-
tila 2010). According to these authors, the rac-
coon dog specialises more on plants and small
mammals and the badger more on inverte-
brates. e preferred habitats of these species
also dier: raccoon dogs prefer meadows, gar-
dens and open woodland with tall and abun-
dant undergrowth, whereas badgers prefer
pine, deciduous and mixed forests with thick
canopy but sparse undergrowth. Kauhala &
Auttila (2010) concluded that the two species
have dierent habitat preferences and there-
fore can coexist in an area. In Poland, facili-
tative interactions between badgers and rac-
coon dog contributed positively to the invasion
success of the second (Kowalczyk et al. 2008).
Raccoon dogs used badger setts as shelter from
cold weather and to avoid predation. ese two
species could even overwinter in the same sett,
using dierent parts. Badger densities did not
show any decline as a consequence of this inter-
action. On the other hand, sett sharing could
be dangerous for badgers because of trans-
mission of diseases and exchange of parasites
(Kauhala & Holmala 2006). Overall, badgers
and raccoon dogs apparently have adapted to
coexist and make use of the available resources
with minimal competition, by using dierent
resources in the same habitat (Jedrzejewska &
Jedrzejewski 1998).
In Mediterranean ecosystems the Iberian
lynx (Lynx pardinus) and the badger are sym-
patric (Martín-Franquelo et al. 1995). ese
species have a similar size, are active during
twilight (Palomares & Delibes 1997, Macdon-
ald 2009) and prey on rabbits as a major food
source (Delibes & Calderón 1979, Martín-
Franquelo et al. 1995). erefore, the niches of
these two species may overlap during the year.
However, they seem to be able to pacically
cohabit the same area by selecting dierent
prey size and adopting slightly dierent activ-
ity patterns: lynxes catch larger rabbits and are
most active at sunrise and dusk whereas badg-
ers prey on small rabbits and are mainly noc-
turnal (Fedriani et al. 1999). Although badgers
are also reported to be crepuscular (Macdon-
ald 1984, Kowalczyc et al. 2003, Do Linh San
et al. 2010), they seem to adjust their habits in
order to cohabit peacefully with the lynx.
Other Carnivora, such as the golden jackal
(Canis aureus), stone marten (Martes foina)
and even otters (Lutra lutra), have only been
found to compete with badgers to some extent
in unusual situations, i.e. in strongly reduced
badger populations as in Bulgaria (Griths &
omas 1993).
Diseases and parasites
Badgers are highly susceptible to Mycobac-
terium bovis infection, the cause of bovine
tuberculosis (Gormley & Costello 2003). is
is a major mortality factor of badgers in Ire-
land and the United Kingdom (Olea-Popelka
et al. 2003). It is also present in Spain, France
and Switzerland (Gortazar et al 2011, Payne
et al. 2013, Schoening et al. 2013). Some stud-
ies indicate that badgers are a reservoir of cat-
tle infection in south-west England, Wales and
Lutra_57_2_Text_v3.indd 98 06/02/2015 21:27
Piza Roca et al. / Lutra 2014 57 (2): 87-109 99
Ireland (e.g. Krebs et al. 1997). However, pub-
lished results are contradictive about whether
culling badgers is an eective measure to
reduce or eliminate bovine tuberculosis in cat-
tle or whether it is even counterproductive, as
badger dispersal may increase as a result of it
(Gallagher & Clion-Hadley 2000, Donnelly
et al. 2003, Grin et al. 2005, Woodroe et al.
2006). Still, culling of badgers has been per-
formed as a measure against bovine tubercu-
losis all around Europe, severely threatening
badger populations (Griths & omas 1993),
especially in the United Kingdom (Dolan
1993).
e outbreak of rabies from the 1950s on,
together with the subsequent attempts to con-
trol rabies, was a major reason for the decline
of badger populations all around Europe in the
20th century (Griths & omas 1993, Smith
2002). Although the red fox is the main res-
ervoir for rabies in Europe, badgers were also
infected in many European countries (WHO
1978–2013). Rabies infection can potentially
reduce badgers’ population densities by 90%
(Schwierz & Wachendörfer 1981). In the United
Kingdom, badgers have contributed to rabies
outbreak (Macdonald 1995, Morgan 1995) and
it is not clear whether vaccination is an eec-
Figure 5. Illustration of four dierent landscapes that the European badger inhabits across its geographical range,
each showing the elements that the badger preferably selects from. e very dierent landscape compositions illus-
trate the adaptability of the species. Upper-le, the landscape in the United Kingdom is represented, with a great
proportion of grassland rich in earthworms, arable land, woodland, hedgerows and other small landscape elements
providing cover, as well as some terrain heterogeneity. Upper-right, the landscape in the Netherlands where badgers
occur is illustrated, with a big proportion of woodland, terrain heterogeneity and cover elements, sandy and dry soils
and little presence of elds and grassland. On the lower right side a Polish landscape is represented, with a dominant
presence of woodland, a minor presence of grassland and arable elds, and including some rocks and other small ele-
ments for cover. Finally, on the lower le side the landscape in Spain is illustrated, poor in woodland, with rocks and
scrubs providing shelter and rabbits as an important food source. Illustration: Ed Hazebroek.
Lutra_57_2_Text_v3.indd 99 06/02/2015 21:27
100 Piza Roca et al. / Lutra 2014 57 (2): 87-109
tive measure (Smith 2002). However, the num-
ber of infected animals has decreased signi-
cantly during the last decade, constituting only
0.5% of rabies cases in Europe from 2000 on
(WHO 1978–2013) and may thus have only a
limited eect on badger populations.
Other diseases to which badgers have been
reported to be vulnerable include mustelid
herpesvirus-1, canine distemper, arterioscle-
rosis, pneumonia, pleurisy, nephritis, enteri-
tis, polyarthritis and lymphosarcoma (Harris
& Yalden 2008). However, these diseases are
of much lower concern compared to bovine
tuberculosis or rabies and are therefore much
less studied. Although these diseases may
aect mortality of badgers, the impact on
badger populations is lower.
Internal parasites common in badgers are
trematodes, nematodes and several species of
tapeworms (Harris & Yalden 2008). Cubs are
also very susceptible to a coccidian parasite
(Eimeria melis) (Anwar et al. 2000). Potential
ectoparasites include eas (Paraceras melis -
badger ea, Chaetopsylla trichosa and Pulex
irritans), lice (Trichodectes melis) and ticks
(Ixodes ricinus, I. canisuga, I. hexagonus, I.
reduvius and I. melicula). Badgers also suer
from mange (Harris & Yalden 2008). To coun-
teract this problem, badgers spend much time
practising self and social grooming (Stewart
& Macdonald 2003). Parasites are of general
low concern, because they do not have an
important economic impact and the power of
spreading is lower, and they also have a much
lower impact on badger populations than
bovine tuberculosis and rabies.
Discussion and conclusions
Main habitat characteristics
e reviewed literature shows that a variety
of factors aect the distribution and spatial
population dynamics of the European badger
around Europe (see also table 1): e.g. climate
and terrain characteristics such as soil type,
slope, heterogeneity and cover. Habitat com-
position, the presence of woodland, grass-
land and crops - such as maize, wheat and
barley - and food availability are also of great
importance (Kruuk 1989, Feore & Montgom-
ery 1999). Built-up areas and roads negatively
inuence badger distribution through habitat
fragmentation, while roads are also an impor-
tant cause of mortality. Hunting, although
forbidden or strictly regulated nowadays in
most countries, is still allowed in some coun-
tries and, together with poaching, contributes
to badger mortality. Biotic interactions such
as interspecic competition are also explain-
ing badger territory expansion. Finally, dis-
eases may aect badger occurrence and den-
sities.
However, the degree of inuence of dierent
factors varies greatly. According to the col-
lected ndings, the main factors enhancing
badger distribution and population densities
are those that favour sett building and food
availability. On the one hand, sett construc-
tion is mostly promoted by factors providing
shelter and facilitating sett excavation, that
is the presence of woodland and other cover
features such as hedgerows and shrubs, ter-
rain heterogeneity, soils that are not too wet
or dicult to dig, and distance to urban areas
and roads. On the other hand, food availa-
bility is enhanced mostly by the presence of
grassland, crops and woodland. It seems that
it is a balance between these two needs which
nally determines the habitat preferences
of the European badger. e optimal habi-
tat composition is given when both are sup-
ported. Several studies conrm the combined
importance of food and sett site availability
(e.g. Woodroe & Macdonald 1993, Rosalino
et al. 2005). More specically, Rosalino et al.
(2005) studied the relationship between food
patches availability and suitable sett site avail-
ability and concluded that the presence of
both factors was required by badgers and that
either one or the other could act as a limiting
factor for badger colonisation and density.
e European badger is, according to all the
Lutra_57_2_Text_v3.indd 100 06/02/2015 21:27
Piza Roca et al. / Lutra 2014 57 (2): 87-109 101
sources we studied, very adaptive and able to
make use of dierent environmental factors
(gure 5). Badgers can nd shelter in forests,
in human-made hedgerows, in a heteroge-
neous rocky area, etc. eir diet is also very
exible, although earthworms are preferably
taken when it is available in the habitat. If
eart hworms are not abundant enough to cover
their needs, this opportunistic omnivore can
feed from other sources such as cereals, small
animals and plants, wild nuts and fruits, or
even anthropogenic food near urban areas.
e home range of the badger is also adapta-
ble to dierent environments. As contractors,
badgers will always establish a territory that
has a minimum land size to cover their needs.
But if the environmental factors for protec-
tion, shelter and feeding such as woody areas
and food patches are too far from each other,
badgers will increase the size of their territory
in order to include the necessary environ-
mental features. On the other hand, if food
availability is low, they will also increase their
home range and travel long distances daily to
nd the needed food. In conclusion, the Euro-
pean badger is a very opportunistic and adap-
tive animal, a fact that is continuously being
rearmed by ecological studies performed on
this species (e.g. Remonti et al. 2006). How-
ever, all over their distribution range, badgers
preferably use certain environmental items
and terrain characteristics. us, all around
Europe the badger shows a dened general
pattern in niche characteristics, indicating a
common realised niche.
Regional variation and differentiation
Although the realised niche might give an
approximation of the fundamental niche of
the species, these may not always overlap.
e species’ ecological fundamental niche
involves all conditions allowing long-term
survival of the species in an area, while the
realised niche is the portion of the funda-
mental niche that the species actually occu-
pies (Hutchinson 1957). e realised niche is
smaller than the potential ecological range
due to various constraints, notably human
pressure, biotic interactions and geographic
barriers. erefore, it is dicult to estimate
the entire fundamental niche of the badger, as
only some components of the full ecological
potential or fundamental niche are expressed
depending on the environmental conditions.
Apart from habitat composition, we have
seen how many other factors can inuence,
to a larger or smaller extent, the population
dynamics of the badger, namely climate,
anthropogenic impact through urban infra-
structure, agriculture, roads and hunting,
as well as diseases such as bovine tuberculo-
sis and rabies. e magnitude of these inu-
ences varies depending on the region, the
environmental and landscape characteristics,
abiotic and biotic composition, history, the
degree of badger protection, etc. erefore,
the most important factors aecting the dis-
tribution and density of the badger will dier
depending on the study area. Also, environ-
mental drivers do not equally aect the dier-
ent elements of spatial population dynamics.
For example, hunting may aect population
size and density, while roads may also aect
colonisation and migration. us, depending
on the region and the focus of the study, the
appreciation of the importance of environ-
mental factors on the distribution and density
of badgers may vary.
Reflections on historical ecology and
niche evolution
Given the great adaptability of the badger, it
is not surprising that they can easily adapt to
new human modied landscapes, and even
benet from anthropogenic transformation
of the landscape (e.g. Huck et al. 2008, Do
Linh San et al. 2011). Being such an adap-
tive species, badgers modify their realised
niche according to the environmental cir-
cumstances of the moment. Consequently,
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102 Piza Roca et al. / Lutra 2014 57 (2): 87-109
the actual niche of the badger may be better
understood by looking at the historical ecol-
ogy of the species, which explains habitat pref-
erences by exible opportunism and adapta-
tion rather than by intrinsic xed preferences.
Having this in mind, some reections can be
made on the historical niche evolution of the
species. Using the United Kingdom as a case
example, the preference for woodland could
be partly explained by the human prosecution
of the badger in agricultural zones due to crop
damage (Moore et al. 1999), rather than by
actual preference for this habitat. Moreover,
sometimes agricultural elds are surrounded
by electrical fences to prevent badger access
(Poole et al. 2002), crops are treated with
repellent to inhibit badger feeding (Baker et.
al. 2005) and farmers even illegally cull badg-
ers to avoid crop damage (Enticott 2011). e
badgers would then select woodland not for
its better conditions compared to agricultural
land but for its less signicant human nega-
tive intervention and the impediment to use
the agricultural elds. Also, the preference for
sloped areas could be partly explained by the
distribution of habitats in relation to terrain
characteristics. Agricultural activities are
preferably performed in at land, while the
more sloped land is le out of deforestation.
erefore, the badgers’ preference for slopes
can be the consequence of an artefact, i.e.
their apparently preference for woodland (or
forced avoidance of agricultural land). Like-
wise, badger setts in urban areas are also rea-
son for human conict (Davison et al. 2011)
and the exclusion of badger from these setts
could explain the avoidance of urban areas in
the badger’s distribution. In conclusion, we
must be careful when drawing conclusions
about habitat requirements and preferences
by looking only at the actual distribution and
density of the species. We should also relate
the niche evolution to the historical ecology
and environmental transformation that the
species has experienced to fully understand
its habitat relationships. is may have impli-
cations for conservation and management,
as the most important management strategy
might not be the availability of the environ-
mental items selected by the European badger
in recent times, but its actual feasibility of
using the dierent habitat elements.
Recommendations
is review provides an overview of factors
aecting the European badger’s distribution
and density. is knowledge can be highly use-
ful for future ecological research on this spe-
cies. e main factors inuencing the badger’s
spatial dynamics are those favouring both sett
location and food availability. erefore, mul-
tiple environmental factors contributing to
these two requirements interact to favour the
badger’s presence and numbers in a certain
area. Sett location requirements are most oen
enhanced by coverage and protection of wood-
land and other small elements, such as shrubs
and hedgerows, and food availability is most
oen higher in grassland. However, depend-
ing on the study area this might vary and other
elements may gain importance, such as suita-
ble soil for sett building, specic crops, human
inuence, diseases, etc. Consequently, prior to
every ecological study in which factors aect-
ing the distribution and density of the Euro-
pean badger play an important role, a choice
of the, potentially, most relevant factors has to
be made carefully according to the study area
characteristics.
Second, this literature review might also
have implications for management. e Euro-
pean badger’s great ecological exibility could
mean that the potential success rate of reha-
bilitation and reintroduction programs should
be relatively high. is may encourage policy
makers to take action when needed. However,
natural colonisation is a slow process in badg-
ers (Reason et al. 1993) and therefore providing
articial setts and translocation of displaced
badger social groups may facilitate and acceler-
ate badger expansion when desired.
ere is a clear bias for research on the badger
Lutra_57_2_Text_v3.indd 102 06/02/2015 21:27
Piza Roca et al. / Lutra 2014 57 (2): 87-109 103
in the United Kingdom, as 50% of the relevant
data on factors aecting t he distribution and den-
sity of badgers where performed in this European
region. However, the environmental conditions
are dierent throughout European regions and
therefore the ndings in the United Kingdom
may not be applicable to other European coun-
tries. We encourage research on badger ecology
in the countries where this is scarce, such as Italy,
France, Belgium or Sweden.
Although extensive knowledge on factors
aecting the distribution and densities of the
European badger is available, little research has
been conducted on the specic eects of environ-
mental factors on life-cycle components, such as
age-specic survival and reproduction rates. Sev-
eral studies focused on the inuence of environ-
mental factors, such as roads or climatic condi-
tions on badger survival (e.g. Clarke et al. 1998,
Macdonald et al. 2010), but these do not include
the eects on other life-cycle components. In fact,
very little is known about the inuence of envi-
ronmental factors on other life-cycle characteris-
tics such as group dynamics, reproduction or dis-
persal. Future research on badger ecology should
try to answer the question of how and how much
all components of the life cycle are aected and
what the integrated eect is on the spatial popula-
tion dynamics of the badgers.
Acknowledgements: We are grateful for nancial sup-
port from the Netherlands Organization for Scientic
Research (NWO-meerwaarde grant 850.11.001 to EJ).
We are thankful for constructive comments by two
anonymous reviewers.
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Samenvatting
Verspreiding en dichtheid van de das
(Meles meles): een literatuuronderzoek
naar sturende omgevingsfactoren
Deze literatuurstudie gaat over de milieufac-
toren die het voorkomen en de dichtheid van
dassen bepalen. De geraadpleegde literatuur,
uit de periode 1970-heden, laat zien dat de das
zich aan verschillende situaties kan aanpas-
sen. Gezien over het hele Europese versprei-
dingsgebied van dassen wordt een algemeen
patroon zichtbaar van geprefereerde omge-
vings- en milieufactoren. De meest bepalende
factoren blijken factoren te zijn die het voed-
selaanbod beïnvloeden en de geschiktheid
van de bodem voor het graven van een burcht.
Meer speciek gaat het om een geschikt
bodemtype om gemakkelijk in te kunnen
graven, kleinschalige heterogeniteit van het
landschap voor dekking en de hoeveelheid
bos en grasland met veel regenwormen. Hoe
belangrijk specieke factoren zijn voor de das
is per gebied verschillend, waardoor de aan-
wezigheid en de dichtheid van dassen wordt
bepaald door de unieke samenstelling van
een gebied, land of regio. We geven aan hoe
de kennis over omgevings- en milieufacto-
ren gebruikt kan worden voor ruimtelijke
modelstudies, natuurbeheer en toekomstig-
onderzoek naar habitatgeschiktheid en dicht-
heid van dassenpopulaties. Desondanks is
meer onderzoek nodig om beter en in detail
te kunnen begrijpen op welke wijze de fases
van de levenscyclus van de das worden beïn-
vloed door de gevonden specieke factoren en
wat het (cumulatieve) eect daarvan is op de
(ruimtelijke) populatiedynamiek van dassen.
Received: 27 October 2014
Accepted: 13 December 2014
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