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Military training areas facilitate the recolonization of wolves in Germany

  • LUPUS German Institute for Wolf Monitoring and Research
  • Senckenberg Research Institute and Natural History Museum Frankfurt


Wolves (Canis lupus) are currently showing a remarkable comeback in the highly fragmented cultural landscapes of Germany. We here show that wolf numbers increased exponentially between 2000 and 2015 with an annual increase of about 36%. We demonstrate that the first territories in each newly colonized region were established over long distances from the nearest known reproducing pack on active military training areas (MTAs). We show that MTAs, rather than protected areas, served as stepping‐stones for the recolonization of Germany facilitating subsequent spreading of wolf territories in the surrounding landscape. We did not find any significant difference between MTAs and protected areas with regard to habitat. One possible reason for the importance of MTAs may be their lower anthropogenic mortality rates compared to protected and other areas. To our knowledge, this is the first documented case where MTAs facilitate the recolonization of an endangered species across large areas.
Received: 20 July 2018 Revised: 12 October 2018 Accepted: 27 January 2019
DOI: 10.1111/conl.12635
Military training areas facilitate the recolonization of wolves
in Germany
Ilka Reinhardt1,2 Gesa Kluth1Carsten Nowak3Claudia A. Szentiks4Oliver Krone4
Hermann Ansorge5Thomas Mueller2,6
1Lupus – German Institute for Wolf
Monitoring and Research, Spreewitz,
2Goethe-University Frankfurt am Main,
Department of Biological Sciences, Germany
3Senckenberg Research Institute and Natural
History Museum Frankfurt, Gelnhausen,
4Leibniz-Institute for Zoo and Wildlife
Research, Department of Wildlife Diseases,
Berlin, Germany
5Senckenberg Museum of Natural History,
Görlitz, Germany
6Senckenberg Biodiversity and Climate
Research Centre, Frankfurt am Main,
Ilka Reinhardt, LUPUS – German Institute
for Wolf Monitoring and Research, Dorfaue 9,
D-02979 Spreewitz, Germany.
Funding information
Robert Bosch Stiftung; Regina Bauer Stiftung
Wolves (Canis lupus) are currently showing a remarkable comeback in the highly frag-
mented cultural landscapes of Germany. We here show that wolf numbers increased
exponentially between 2000 and 2015 with an annual increase of about 36%. We
demonstrate that the first territories in each newly colonized region were established
over long distances from the nearest known reproducing pack on active military
training areas (MTAs). We show that MTAs, rather than protected areas, served as
stepping-stones for the recolonization of Germany facilitating subsequent spreading
of wolf territories in the surrounding landscape. We did not find any significant differ-
ence between MTAs and protected areas with regard to habitat. One possible reason
for the importance of MTAs may be their lower anthropogenic mortality rates com-
pared to protected and other areas. To our knowledge, this is the first documented case
where MTAs facilitate the recolonization of an endangered species across large areas.
Canis lupus, large carnivores, population growth, protected areas, recolonization
After their all-time low in the 1960s (Boitani, 2003), wolves
(Canis lupus L.) currently show a remarkable return in cen-
tral and western Europe (Chapron et al., 2014). The most
important reason fostering the recovery of large carnivores
in Europe was changes in legislation improving their protec-
tion status that were put into place in the 1980s and 1990s
(Bern Convention and the Habitat directive). At the same
time, ungulate populations have been increasing in many parts
of Europe (Boitani & Ciucci, 2009; Linnell & Zachos, 2010)
and public attitudes toward wildlife conservation, including
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original
work is properly cited.
© 2019 The Authors. Conservation Letters published by Wiley Periodicals, Inc.
large carnivores, have shifted to the positive, providing favor-
able conditions for their return (Boitani & Ciucci, 2009).
This is also true for Germany where wolves were eradicated
in the 19th century. In the German Democratic Republic, state
policy prevented any resettlement of the wolf; while in the
Federal Republic of Germany, wolves had been strictly pro-
tected since 1980, albeit absent. Only after the German reuni-
fication in 1990, the wolf became strictly protected through-
out the country (Reinhardt & Kluth, 2007; Reinhardt, Kluth,
Nowak, & Mysłajek, 2013). In consequence, wolves emigrat-
ing from Poland have recolonized Germany since the late
1990s (Reinhardt & Kluth, 2007; Reinhardt et al., 2013). The
Conservation Letters. 2019;e12635. 1of7
FIGURE 1 (a) Exponential increase of reproductive wolf units in Germany from 2000 to 2015. (b) Distribution of all wolf territories
documented in Germany in 2015 (data from DBBW, 2017)
first colonization area was in the Saxony-Brandenburg region
(Figure 1b) close to the Polish border where, in 2000, the
first reproduction of wild wolves was documented (Kluth,
Ansorge, & Gruschwitz, 2002). Since then, a rapid recolo-
nization in Germany led to a population of 47 wolf packs and
21 pairs in 2015/2016 (Dokumentations- und Beratungsstelle
des Bundes zum Thema Wolf [DBBW], 2017). Wolves spread
across the country with wolf territories in 7 out of 16 federal
states in 2015 (Figure 1b). Although wolves are known to be
adaptive in respect to habitat requirements (Boitani, 2000),
they prefer sites where the degree of human disturbances
is low, especially when rearing pups (Kaartinen, Luoto,
& Kojola, 2010; Llaneza, García, Palacios, Sazatornil, &
López-Bao, 2016; Sazatornil et al. 2016). This raises the ques-
tion how the remarkably fast population recovery in a densely
populated and highly fragmented country like Germany was
possible. How important, for example, were protected areas
(PAs), for the recolonization process? Interestingly, the first
wolf territory occurred on an active military training area
(MTA) (Kluth et al., 2002). Subsequently, wolf territories
have been established on different MTAs but also on PAs, and
other lands (Reinhardt & Kluth, 2007; Bundesministerium
für Umwelt Naturschutz Bau und Reaktorsicherheit [BMUB],
2015). We quantified the population growth, analyzed mor-
tality events, and reconstructed the spatial patterns of wolf
recolonization based on existing data from official monitor-
ing reports and a public database. We focused, in particu-
lar, on the role of PAs and MTAs for the recovery of wolves
in Germany. While MTAs are known to be valuable areas
for nature conservation (Aycrigg, Belote, Dietz, Aplet, &
Fischer, 2015; Lindenmayer et al., 2016; Zentelis & Lin-
denmayer 2015; Zentelis, Lindenmayer, Roberts, & Dovers,
2017), their role in facilitating recolonization processes of
large mammals in human dominated landscapes has not been
explored to date. We expected MTAs and PAs, to be important
areas for the recovery of wolves in Germany.
We estimated the population growth of wolves in Germany
since their return in 2000 until 2015 by using the number of
reproductive units (packs and pairs). These data are annually
collected according to the rigorous German national monitor-
ing standards for large carnivores (Reinhardt et al., 2015) and
are yearly published in official reports (Reinhardt et al., 2013;
DBBW, 2017). We then fitted an exponential growth model
to the data and calculated the annual population growth. For
all statistical analyses, we used the programming language R
(R Core Team 2016). For analyzing the spatial spread, we
referred to established wolf territories (of packs, pairs, or sin-
gle resident wolves) that were identified and defined by cri-
teria of the national monitoring standards (Reinhardt et al.,
2015). The location types of the territories were classified
as “protected area,” “military training area,” or “other” (for
details, see the Supporting Information). We used the federal
states of Germany as spatial scale of analysis, because wolf
monitoring is conducted on a state-by-state basis. We then
identified where the first (up to five) territories were located in
each federal state after the initial colonization happened in the
Saxony-Brandenburg border region (Supplemental Figure 1).
In order to exclude short-term occurrences, we only included
territories that were occupied for at least three consecutive
years and still existed when analyzing the data in 2017 (long-
term territories).
To analyze how far wolves moved to establish new territo-
ries, we calculated the minimum dispersal distance for all wolf
territories established between 2000 and 2015 by measuring
the distance between the focal territory and the next known
potential source territory in Germany (i.e., the next pack with
confirmed reproduction in the previous year). Using the min-
imum dispersal distance, we ignored that territory founders
might have immigrated from Poland or other European coun-
tries. However, comparing our distribution data to Poland
(Nowak & Mysłajek 2016) or other countries, the next possi-
ble source territory outside Germany was usually farther away
for newly established wolf territories than the next source
territory within Germany. Our dispersal distances are there-
fore conservative. We log-transformed the dispersal data and
performed linear mixed models with federal state as random
effect to test whether the minimum dispersal distance differed
between territories established on MTAs, PAs, or other areas.
In addition, we mapped the dispersal pattern for the founding
individuals of the first two long-term wolf territories in each
state (Figure 3b), because, for these individuals the natal terri-
tories were available from the German wolf database (DBBW,
For each territory that has been established between 2000
and 2015, we analyzed two key habitat variables that deter-
mine habitat suitability for wolves in neighboring Poland –
forest cover and road density (Jedrzejewski et al., 2008). The
habitat values were extracted from CORINE Land Cover clas-
sification raster data (2012) and the Digital Landscape model
of the German Federal Agency for Cartography and Geodesy
(Bundesamt für Kartographie und Geodäsie, 2018).
Finally, we analyzed wolf mortalities from 2000 to 2015
using the publicly available German wolf database (DBBW,
2017) where the cause of each mortality event is classified
as anthropogenic (traffic or poaching), natural, or unknown.
We included only mortality events found within known wolf
territories (n=92) and assigned each mortality location to
MTA or PA within a 1 km buffer or to other area. For each
of the three location types, we calculated the number of “ter-
ritory years,” that is the cumulative number of years territo-
ries of each location type existed. This allowed us to score the
amount of mortality cases found in each location type rela-
tive to the number of wolf territories and the time they had
been occupied. We used Pearson's chi-squared test with sim-
ulated P-values (based on 2000 replicates) for comparing the
number of mortalities relative to “territory years” among
MTAs, PAs, and other areas.
We found that the increase of wolf reproductive units in Ger-
many from one unit in 2000 to 67 in 2015 followed an expo-
nential growth curve with a 36% increase per year (Figure 1a).
The first three wolf territories were located on an active
MTA in Saxony. The three subsequent territories were
established in their immediate surroundings forming together
a relatively small initial colonization area (Figure 1b and Sup-
plemental Figure 1). In 2007, the first two wolf territories out-
side the initial colonization area were established more than
200 km away on two MTAs (Supplemental Figure 1). In the
following years, more territories were established in new fed-
eral states, long distances from the nearest reproducing pack.
FIGURE 2 Probability of territory establishment on MTAs
relative to their order of establishment (time rank) in each state (n=23
territories from six states). The gray curve represents a logistic model
fit. Note that no territories on protected areas were among the first three
in any state (not shown)
We found the first and second long-term territory in each of
these newly colonized states were always established on active
MTAs (Figure 2 and 3a and 3b). None of the initial territories
were established on PAs or other areas.
Between 2000 and 2015, 16 out of 79 territory establish-
ments took place on active MTAs, 9 on PAs, and 54 in other
areas. For wolves founding a new territory on a MTA, the
average minimum dispersal distance was considerably greater
(128 km, median 165 km) than for wolves that established
their territories on PAs (64 km, median 31 km, 𝛽=–0.356.27,
P<0.05) or on other areas (38 km, median 20 km, 𝛽=–0.596,
P<0.001; Figure 4).
Some of these initial recolonizing events in new federal
states far away from the next source pack served as stepping-
stones allowing subsequent colonization in the surroundings
areas (Figure 1 and Supplemental Figure 1). In 2015, 13 out
of 21 MTAs (62%) with a minimum size of 30 km2were
occupied by wolves, but only 8 out of the 55 (14%) PAs of
the same size class were occupied.
We found no significant difference between MTA territo-
ries and PA territories in the two key habitat variables forest
cover (mean MTA: 52%, mean PA: 50%, tvalue =–0.263,
P>0.1) and road density (mean MTA: 0.48 km km−2, mean
PA: 0.59 km km−2,tvalue =0.601, P>0.1; Supplemen-
tal Figure 2). Likewise, the amount of forest cover was not
different between MTA territories and other territories (mean
other areas: 47%, tvalue =–1.001, P>0.1). The only sig-
nificant habitat difference we found was in the road density
FIGURE 3 Establishment patterns of the first two permanent territories per federal state from 2000 until 2015. (a) The first territories were
always established on MTAs. (b) Origin of the founder animals of the first long-term territories. Red arrows: females, blue arrows: males. Dotted
lines: individual immigrated from Poland. Solid lines: Individual dispersed from known pack in Germany (for Brandenburg only territories outside
the initial colonization area were considered in Figures 3a and 3b)
FIGURE 4 Median and range of minimum dispersal distances
for wolves establishing territories on MTAs, PAs, and other areas
between MTA territories and other territories (mean other
areas 0.61 km km−2,tvalue =2.385, P<0.05).
Anthropogenic mortality was the prevailing mortality
cause accounting for 80% (n=74) of recorded deaths within
territories. Anthropogenic fatalities were lower on MTAs
compared to other areas (relative to the total territory years
in these location types [see Table 1], chi-square =9.65,
P<0.01). There was no difference between anthropogenic
fatalities on PAs compared to other areas (relative to the
total territory years in these location types [see Table 1], chi-
square =0.59, P=0.46). Mortality due to poaching was
higher on PAs than on MTAs, although the sample size was
relatively low (Table 1; chi-square =13.41, P<0.001).
This is the first study that examines the role of active MTAs in
facilitating the recolonization of a previously extirpated large
carnivore in a highly human modified landscape. The rapid
rate of population increase and range expansion of wolves
in Germany was facilitated by the presence of MTAs. These
sites, rather than PAs, acted as stepping-stones promoting the
recolonization of new areas far away from the next source
This form of jump expansion with initially large gaps
between wolf territories was previously reported from other
regions (Nowak & Mysłajek 2016; Wabakken, Sand, Liberg,
& Bjärvall, 2001; Wydeven, Schultz, & Thiel,, 1995). Unique
about the expansion in Germany is that MTAs were exclu-
sively used as stepping-stones. Once wolves established ter-
ritories and bred on MTAs a subsequent diffusion like range
expansion around these initial colonization areas could also be
observed. There are three potential explanations for the initial
preference of active MTAs: (1) habitat preference, (2) natal
habitat preference, or (3) mortality risk.
1. MTAs are known to play an important role for con-
servation as they harbor disproportionally high
numbers of threatened and endangered species
(Stein, Scott, & Benton, 2008; Warren et al., 2007).
Biodiversity on these sites is often high even compared to
national parks (Arimoro et al., 2017; Aycrigg et al., 2015;
Flather, Joyce, & Bloomgarden, 1994; Groves et al., 2000;
Stein et al., 2008; Warren et al., 2007). The conservation
effect of MTAs is often linked to an artificially maintained
patchiness favoring species richness in plants (Jentsch,
Friedrich, Steinlein, Beyschlag, & Nezadal, 2009; Molino
& Sabatier 2001), invertebrates (Cizek et al., 2013; War-
ren & Büttner, 2008), and birds (Gazenbeek, 2005). In
TABLE 1 Number of territory years (cumulative number of years territories were occupied) and the number of wolves found dead within
Location Territory years Mortality total Mortality traffic Mortality poaching
Military training areas 103 17 (0.16) 13 (0.13) 0 (0.0)
Protected areas 42 15 (0.36) 5 (0.12) 6 (0.14)
Other areas 142 60 (0.42) 44 (0.31) 6 (0.04)
Locations where carcasses were found were classified per location type as military training area, protected area, or other area. Traffic and poaching mortality sums up to
anthropogenic mortality. Total mortality includes anthropogenic, natural, and unknown (the latter two not shown). The number in brackets denotes mortality cases found
per territory year.
providing and maintaining rare habitat conditions, MTAs
may serve as refugee areas especially for some habitat
specialists (Jentsch et al., 2009; Warren & Büttner, 2008).
Wolves, however, are habitat generalists that are known
to adapt to a wide variety of ecological conditions (Fritts,
Stephenson, Hayes, & Boitani, 2003). Among Europe's
large carnivore species, wolves are the most successful
in adapting to human-dominated landscapes (Chapron
et al., 2014). This habitat adaptability of wolves is also
evident in Germany where after the initial establish-
ment of territories on MTAs subsequent territories were
established in other areas. In addition, we did not find
significant differences in key habitat variables between
MTA territories and PA territories. Forest cover did not
differ between MTA territories and PA territories nor did
road density. If forests would have played a key role in
the colonization process, we would have expected the
large forest complexes of north-east Germany close to
the Polish border and source population (Czarnomska,
Borowik, Niedziałkowska, Stronen, & Nowak, 2013) to
be recolonized first (Reinhardt & Kluth, 2007; BMUB,
2015). However, wolves did not settle there until 2015
(Supplemental Figure 1). Differences in habitat type
or road densities thus cannot explain the preference of
MTAs. Because densities of wild ungulates, the main
prey of wolves in Germany (Wagner, Holzapfel, Kluth,
Reinhardt, & Ansorge, 2012), are high in all areas settled
by wolves to date (Reinhardt & Kluth, 2007), it is also
implausible that different prey densities could serve as an
explanation. Overall, it is unlikely that habitat suitability
alone was the primary driver for the strong initial selection
for MTAs by wolves.
2. The preference for MTAs may be partly explained with
natal habitat preference. Natal habitat preference has been
shown in a variety of species where dispersing animals
tend to choose habitat types similar to those where they
have been raised (overview in Stamps & Davis, 2006).
Indeed, seven out of eight wolves with known natal ter-
ritories (i.e., wolves born in Germany) that settled on
MTAs have also been raised on MTAs (Figure 3b). For
the wolves emigrating from Poland, the natal territories
remain unknown. We believe it is unlikely that most of
these wolves were raised on MTAs, because MTAs did not
play a critical role during wolf recolonization in Western
Poland (Nowak & Mysłajek 2016; Nowak et al., 2017).
While it is possible that natal habitat preferences play
at least some role in the colonization process, additional
research would be needed for conclusive evidence.
3. The initial preference for active MTAs may at least partly
be linked to the lower level of anthropogenic mortality
on MTAs compared to other areas, including PAs. PAs
did not show a lower mortality rate when compared to
other areas. Traffic incidents are relatively low on MTAs
because of the low frequency of public roads, but this
is also true for PAs. The second component of anthro-
pogenic mortality is poaching. Poaching potentially plays
a greater role than we estimate here because most poach-
ing events remain undetected (cryptic poaching) (Liberg
et al., 2011). One key difference between MTAs and other
areas, including PAs, is the hunting regime. In Germany,
hunting on MTAs is supervised by federal authorities and
is managed across large areas, whereas PAs and other areas
usually are divided in private hunting grounds with a min-
imum size as small as 75–150 ha. This may lead to sit-
uations where a wolf pack shares its territory with more
than 100 hunters which, in turn, make these territories
more vulnerable to poaching even if most hunters do not
poach. For many PAs, the hunting regime often follows the
same small-scale approach because landownership is often
fragmented, including private lands (with the exception of
national parks). Therefore, opinions and attitudes of land
owners and hunters on protected and other areas may differ
considerably leaving more opportunity for illegal killings
than on strictly and uniformly managed MTAs. This may
explain why we found lower poaching rates on MTAs ver-
sus PAs. We believe it unlikely that our findings on low
mortality on MTAs are a result of lower detectability of
carcasses in these areas. Although these sites are closed
for the public, they are intensely used by armed forces and
forestry and subject to a wide range of environmental mon-
itoring programs.
Overall, for the rapid recolonization of wolves in Germany
MTAs uniquely served as stepping-stones despite wolves
settling on MTAs had to disperse longer distances dur-
ing the early years of population recovery as compared to
wolves that settled near their natal territories. A contributing
factor seems to have been the lower mortality rate on MTAs.
At very low population densities, anthropogenic mortality
may have had an additive effect on the wolf population and
may have been simply too high outside MTAs ultimately
shaping this unique colonization pattern. A similar additive
effect of anthropogenic mortality at low population density
has been shown for the red wolf (C. rufus) (Sparkman, Waits,
& Murray, 2011). At today's population densities however,
the anthropogenic mortality rate seems to be less critical and
rather compensatory allowing for a robust population growth.
4.1 Management implications
MTAs are known to be important conservation areas (Lin-
denmayer et al., 2016; Zentelis et al., 2017), but their bene-
ficial effect on large mammalian species at larger scales has
received little attention to date. The beneficial effect may
extend well beyond the wolf example of this study (Arimoro
et al., 2017; Zentelis & Lindenmayer 2015). Though MTAs
are under the influence of human activity and disturbance,
they tend to be less fragmented than other areas (Ibisch et al.,
2016) and serve as refugia in highly human-dominated land-
scapes. MTAs in Germany are the largest land use category
with a unified management of the federal government. The
hunting regime on MTAs is homogeneous over a large area
and may provide less opportunity for poaching.
We conclude that MTAs especially in highly fragmented
Europe are key areas for large carnivore conservation and
make a substantial contribution to conservation outside the
formal protected area network (Lindenmayer et al., 2016).
When these areas are taken out of military use, particular
attention should be paid on how to maintain their function
as refugia for species conservation (Cizek et al., 2013). The
listing of large parts of current and former MTAs as Natura
2000 areas is a first step to preserve their conservation func-
tion. However, we recommend the strict hunting management
for MTAs should continue after the sites become inactive.
This study was supported by the Regina Bauer Stiftung (Ger-
many) and the Robert Bosch Foundation. We thank the Fed-
eral Forest Agency for their support. We thank Bob Hayes and
four anonymous reviewers for improving the manuscript.
IR and TM conceptualized the paper, IR analyzed the data,
and all authors contributed to the writing of the manuscript.
The authors declare no conflict of interests.
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... Following severe historic persecution leading to the absence of large carnivores in many areas of Central Europe during most of the nineteenth and mid-twentieth century, large carnivore populations have increased over recent decades (Chapron et al., 2014). The gray wolf (Canis lupus) has received legal protection in the 1980s in most European countries, and has since increased its geographic range (Nowak and Myslajek, 2016;Reinhardt et al., 2019). In general, wolves can persist in human-modified landscapes as long as human tolerance and policy are favorable (Boitani and Linnell, 2015;Reinhardt et al., 2019). ...
... The gray wolf (Canis lupus) has received legal protection in the 1980s in most European countries, and has since increased its geographic range (Nowak and Myslajek, 2016;Reinhardt et al., 2019). In general, wolves can persist in human-modified landscapes as long as human tolerance and policy are favorable (Boitani and Linnell, 2015;Reinhardt et al., 2019). Apart from legal protection and supportive public opinion, the main factors for large carnivore recovery (Chapron et al., 2014), habitat suitability for wolves has increased over the last two decades in some European areas, including Central and Northern Europe, which is correlated with decreasing human population density and increasing forest cover (Cimatti et al., 2021). ...
... Hence, to reduce conflict levels, keeping damage to livestock at low levels is important (Bautista et al., 2019). To do so, it is crucial to gather knowledge on wolf distribution and habitat use (Reinhardt et al., 2019;Cimatti et al., 2021), impact of livestock density on colonization patterns, as well as identifying spatial centers of livestock predation. Moreover, it is important to evaluate existing livestock protection measures (Eklund et al., 2017). ...
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Attacks by large predators on livestock are an important driver of conflicts. Consequently, knowledge about where predators occur, where livestock depredation takes place and what factors influence it will aid the mitigation of stakeholder conflicts. Following legal protection, wolves (Canis lupus) in Central Europe are recently spreading to areas dominated by agriculture, bringing them in closer contact with livestock. Here, we analyzed habitat selection and livestock depredation rates of 43 wolves identified by genotyping on the Jutland peninsula, consisting of mainland Denmark and the northernmost German federal state Schleswig-Holstein. Occupancy by resident wolves correlated positively with forest and other non-forested semi-natural land cover (habitat for natural ungulate prey), whereas occupancy by non-resident wolves correlated with increasing forest cover and sheep density. The latter effect likely reflected increased sampling probability of highly mobile dispersers killing livestock. We recorded 565 livestock depredation events (85 in Denmark and 480 in Schleswig-Holstein), of which 42% (55 in DK and 185 in SH) could be assigned to 27 individual wolves based on DNA evidence. Livestock (mostly sheep) were killed by wolves in 16% of the study area. Our results indicate that wolves mostly killed livestock as a context-dependent response, i.e., being dispersers in agricultural areas with low availability of wild ungulate prey and high livestock densities, and not because of behavioral preferences for sheep. Moreover, the livestock depredation was lower in areas with livestock protection measures (implemented in areas with established pairs/packs). We conclude that while wolf attacks on livestock in established wolf territories generally can be reduced through improvement of fences, livestock depredation by non-resident wolves in agricultural areas constitutes a bigger challenge. Albeit technically possible, the economic costs of implementing predator-proof fences and other preventive measures in such pastoral areas infrequently visited by wolves will be considerable. Experiences so far further indicate that lethal removal of identified "problem wolves" may be inefficient in practice.
... Recently, the population size was estimated at ca. 1900 individuals (Species Report, 2019) and continues to grow. Polish wolves have also spread to Germany and other western-European countries (Andersen et al., 2015;Reinhardt et al., 2019), where their population is rebuilding. From the perspective of wolf protection in Western Europe, where wolf mortality is high (Chapron et al., 2014;Sonne et al., 2019) it is crucial to maintain vital populations in Central Europe, which provide dispersing individuals to the West. ...
... The biggest number of shot and snared wolves were discovered in Western Poland, where the population recovered during the last 15 years, after dozens of years of absence Mysłajek, 2016, 2017;. It is the Polish part of the Central European wolf population that is categorized as Vulnerable, according to the IUCN Red List of Threatened Species (Boitani, 2018) and is the source of wolves settling in Western Europe (Andersen et al., 2015;Reinhardt et al., 2019;Schley et al., 2021). Also, four from six GPS/GSM collared wolves were illegally shot within the range of the CE wolf population. ...
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In central Europe, wolves Canis lupus prey on wild ungulates - main game species and occasionally kill livestock. The recovery of wolf population across the continent coincides with an increasing incidence of illegal killing, which level remains unknown. We analysed the illegal killing of wolves in Poland, where the species is strictly protected since 1998. We opportunistically collected data on wolves illegally shot and snared from 2002 to 2020, revealing their geographical extent and sex and age structure. Furthermore, we estimated their mortality rate due to illegal shooting on the basis of 16 GPS/GSM collared individuals between 2014 and 2020. We recorded 54 illegally shot and 37 snared wolves. The majority (63.7%) were killed between 2017 and 2020, mostly in Western Poland. The sex structure was similar between shot and snared individuals. In both groups, the wolves over one-year old prevailed, although there were 18 pups among shot wolves. We identified 6 shot and 3 snared breeders. Out of 16 GPS/GSM collared individuals, six were shot giving the mortality rate of 0.33 per year. Simulations revealed that the pooled number of wolves illegally shot in Poland annually, is between 147 and 1134 (99% highest density interval) or 216 and 1000 (95%). In six out of seven cases, in which the person who shot a wolf was eventually sentenced, hunters were responsible. We conclude that the present regulations concerning the prevention of illegal killing, pursuing and punishing the perpetrators of the illegal killing of wolves, require urgent improvements in order to effectively mitigate the problem.
... Large carnivores in Europe predominantly persist outside of protected areas (Chapron et al., 2014), which increases the probability of interactions with humans (Crespin and Simonetti, 2018;Rode et al., 2021). Reducing negative interactions is possible by spatially or temporally segregating human and large carnivore activities Reinhardt et al., 2019). To achieve this separation, large carnivore behavior can be influenced by ensuring favorable habitat conditions in areas away from human settlements, and using physical deterrents to protect livestock, such as fences and guardian dogs . ...
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) Understanding Human–Canid Conflict and Coexistence: Socioeconomic Correlates Underlying Local Attitude and Support Toward the Endangered Dhole (Cuon alpinus) in Bhutan.
... These zones can also have importance as corridors, connecting different patches (Hosseini et al., 2019). Reinhardt et al. (2019) point out that military training areas facilitating the spreading of wolf (Canis lupus) territories in the surrounding landscape may be linked to a low level of anthropogenic mortality. However, there is likely to be no less anthropogenic pressure in borderlands under military control than in demilitarized zones due to the large amount of logistical and military activity caused by the presence of troops. ...
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Geopolitical borderlands are politically sensitive areas and biodiversity hotspots, strictly controlled by the government and military. How to ensure political security, while protecting the biodiversity in borderlands is a problem for ecologists and governments. In this study, the nest site selection of the wild boar Sus scrofa was a case study in the Sino-Russia borderland to understand the survival strategy of wild life under anthropogenic pressure. We investigated (a) how the spatial distribution of anthropogenic pressure and wild boar nests in the borderland and (b) how anthropogenic pressure and the border influence on the wild boars' nest site selection. The Getis-Ord Gi * analysis was used to analyze the distribution patterns of wild boar nest sites and anthropogenic pressures in the borderland, the Structural Equation Models was used to explore the influence of border, roads, settlements, agricultural land, grassland and anthropogenic pressure on wild boars' nest site selection. The results indicated that wild boar nest sites are close to the border, roads and agricultural land and away from settlements and grassland. Regardless of the combination of anthropogenic pressure, wild boars make the most advantageous choice and prefer to be closer to the borderland. We speculated that military control played a vital role in borderlands for animal protection under anthropogenic pressure. Wild boars benefit from the prohibition of anthropogenic persecution due to military control. Compared with existing measures, we suggest a different protection/wildlife management strategy, what we need to do may be to prohibit anthropogenic persecution rather than perform other human interventions to protect animals. However, for a species with trouble potential, we need to base our conservation strategies on the recovery of top predators, and play the community control role of top predators to avoid the occurrence of trouble.
... This delayed onset of depredation might be related to low wolf densities during the initial recolonization phase (Kaartinen et al., 2009;Nowak & Mysłajek, 2016). Additionally, in the initial phases of recovery wolves can inhabit areas distant from human settlements (Nowak et al., 2017;Reinhardt et al., 2019) reducing probability of wolf-livestock encounters. Especially in the Western Lowlands, large compact forests offer optimal habitats for wolves to settle (Gula et al., 2020) away from rural areas. ...
As the wolf Canis lupus populations continue to recover across Europe, livestock depredation becomes increasingly challenging for their effective conservation. We aim to (1) analyze the spatiotemporal variation in wolf attacks on livestock in relation to the landscape structure, livestock species, and the phase of wolf expansion in Poland and (2) discuss the implications for conservation and management in an expanding, protected wolf population. From 2008 to 2018, farmers reported 5499 attacks on livestock with 13,164 killed individuals, and the number of attacks increased 2.7-fold at the country scale. Sheep were among the most frequently killed livestock, and surplus killing (>2 killed individuals) was relatively common in captive deer and sheep depredation. The attacks were patchily distributed; 59% of all kills occurred in municipalities constituting 1% of the country surface. The probability and number of attacks were positively influenced by forest and pasture cover, and by the occurrence of depredation in previous years. Spatial variation and long-term dynamics in livestock depredation by wolves can be attributed to different husbandry practices and phases of wolf recovery in three regions of the country. Our results indicate that accelerating increase in depredation rates during the initial phases of wolf recovery is likely to be followed by stabilization or decrease in attacks. A detailed spatiotemporal analysis of wolf–livestock conflict can help in management decisions in areas with ongoing wolf population recovery.
... Large carnivores are recolonizing former grounds in Europe (Chapron et al., 2014) and North America (Bruskotter and Shelby, 2010), yet carnivore recovery pace and success vary across regions. In central Europe, wolf recovery has been quite fast in Germany (Reinhardt et al., 2019), and wolves even reproduced in Denmark for the first time in~200 years, although poaching may prevent further expansion (Sunde et al., 2021). In northern Europe, the Scandinavian wolf population started its recovery in the 1990s, but nowadays wolves are more numerous in Sweden than in Norway due to differences in policy (Bischof et al., 2020). ...
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Wolf management in Spain is remarkably different at regional scales. South of Douro river, wolves are protected, north of Douro wolves can be hunted, and culling occurs on both sides. After a formal request to include wolves in the Spanish Red List of Threatened Species, wolves have been "listed", but not as a vulnerable species. Recreational hunting will no longer be a wolf management option, while culling is still allowed. We describe the process to raise wolf protection at the state level, and the factors that should be relevant to guide apex-predator management. Restricting lethal control and favoring predator-prey interactions by reducing livestock depredation should be more feasible with an overarching policy that is binding over the whole range of the species in Spain.
... Consequently, the wolf plays a relevant role in the rewilding process that is taking place in many areas where it was extirpated but is now returning. In Europe, the wolf is currently recolonizing historical regions of its distribution range [3,25], for example, in Scandinavia [26], Finland [18,27], Poland [28], Germany [13,29], Denmark [30] and Spain [7,31,32]. This return can result in competition for ungulate game species with hunters and trigger conservation conflicts [18,33,34]. ...
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The ongoing recolonisations of human-transformed environments in Europe by large carnivores like the wolf Canis lupus means that conservation conflicts could re-surface, among other reasons, due to predation on ungulate game species. We estimated the effect of wolves on ungulate species using data on wolf prey selection, kill rates and territory size to build a hypothetical case of future expansion. We extrapolated results on predation from the current wolf distribution in central Sweden and eastern Poland to the eventual wolf recolonisation of southern Sweden. We then calculated the proportion of five ungulate game species killed annually by wolves, and the ratio between the predicted annual predation by wolves given future colonization and the number of ungulates currently harvested by hunters. Results showed that wolf recolonization in southern Sweden would have a minor impact on the estimated population densities of red deer Cervus elaphus, fallow deer Dama dama and wild boar Sus scrofa, but is likely to lead to a significant reduction in human captures of moose Alces alces and roe deer Capreolus capreolus. The current five-ungulate species system in southern Sweden suggests a potential for two to four times higher wolf density than the two-ungulate species system in the northern part of their current distribution. Management and conservation of recolonizing large carnivores require a better understanding of the observed impact on game populations under similar ecological conditions to ameliorate conservation conflicts and achieve a paradigm of coexistence. Integrating these predictions into management is paramount to the current rewilding trend occurring in many areas of Europe or North America.
... Currently, only the city-states of Berlin, Hamburg and Bremen, and the smallest state of Saarland, do not have resident wolves. Wolf numbers in the country grow exponentially, on average by 28% per year, due to population expansion fostered by high mobility, reproductive potential and adaptability of wolves, prey abundance, and the presence of suitable corridors and stepping stones (Reinhardt and Kluth, 2016;Reinhardt et al., 2019Reinhardt et al., , 2021Plaschke et al., 2021). Therefore, it is not surprising that increasing losses of domestic livestock and farmed game species are associated with increasing wolf numbers. ...
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Recovery of predator populations triggers conflicts due to livestock depredation losses, particularly in Germany where the wolf (Canis lupus) population grows exponentially and livestock (especially sheep) losses raise public concerns and motivate the authorities to control wolf numbers. Yet, the effects of wolf numbers and alternative factors, such as abundance of prey and livestock, on livestock losses in this country are not investigated. In this study, we collected and analyzed data on the numbers of reproductive units of wolves (packs and pairs together) as a surrogate of adult wolf numbers, sheep killed by wolves, living sheep, red deer (Cervus elaphus), roe deer (Capreolus capreolus), and wild boar (Sus scrofa) in every German state and year from 2002 to 2019. We applied a negative binomial Generalized Linear Mixed Model (GLMM) to estimate the effects of these predictors on the numbers of sheep killed by wolves. We also examined the relationships between the percentages of killed/living sheep and the numbers of living sheep. Ranking of 63 models based on the Akaike information criterion revealed that sheep losses were determined by state, year, and number of living sheep, not by wolf numbers, at high precision and accuracy. The number of sheep killed by wolves increased consistently by 41% per year and by 30% for every additional 10,000 sheep, mainly in the north where most wolf territories are concentrated. This means that sheep are protected insufficiently and/or ineffectively. The percentages of killed/living sheep consistently increased by 0.02-0.05% per state and year, with the maximum percentage of 0.7%, on a backdrop of decreasing numbers of living sheep. In conclusion, we demonstrate that sheep losses in Germany have been driven by the expansion of the wolf population, not by wolf numbers, and by the number of sheep available. We suggest that Germany's wolf conservation policy should focus on alternative non-lethal interventions, enforcement and standardization of intervention monitoring, and promotion of wolf tolerance rather than on lethal control of wolf population size.
ContextBehavioral adjustments by large carnivores can be a key factor facilitating their coexistence with people in shared landscapes. Landscape composition might be a key factor determining how large carnivores can adapt to occurring alongside humans, yet broad-scale analyses investigating adjustments of habitat use across large gradients of human pressure and landscape composition are lacking.Objectives Here, we investigate adjustments in habitat use by Eurasian lynx ( Lynx lynx ) in response to varying availability of refuge habitats (i.e., forests and rugged terrain) and human landscape modification.Methods We used a large tracking dataset including 434 individuals from seven populations to analyze variation in lynx’ use of refuge habitats and landscape modification at a continental scale.ResultsWe found that lynx use refuge habitats more intensively with increasing landscape modification, selecting forests most strongly in otherwise open landscapes and rugged terrain in mountainous regions. Higher forest availability enabled lynx to place their home ranges in more human-modified landscapes. Human pressure and refuge habitat availability also shaped temporal patterns of lynx habitat use, affecting daytime-nighttime differences in refuge habitat use, as well as females’ association with human-modified habitats during the first months after kittens are born.Conclusions Our findings suggest a remarkable adaptive capacity of lynx towards human pressure and underline the importance of refuge habitats for enabling coexistence between large carnivores and people. More broadly, we highlight that the composition of landscapes determines how large carnivores can adapt to human pressure, and that both factors interact in shaping large carnivore habitat use and distributions at broad scales.
Zusammenfassung Gegenstand und Ziel Seit 2000 siedelt sich der Wolf in Deutschland wieder an. Mit steigenden Wolfszahlen nehmen die durch Wölfe verursachten Nutztierschäden zu, wobei auch Pferde betroffen sind. Ziel der Studie war es, einen Überblick zu dieser Problematik sowie Lösungsansätze zu geben. Material und Methoden Anhand einer Literaturrecherche wurden Daten zur Wolfspopulation in Deutschland, wolfsbedingten Schäden sowie Möglichkeiten und Grenzen des Herdenschutzes beim Pferd eruiert. Eine Online-Befragung von Pferdehaltern/-besitzern diente dazu, die tatsächliche und/oder gefühlte Bedrohung durch Wölfe und die daraus resultierenden Auswirkungen auf die Pferdehaltung zu ermitteln. Ergebnisse Die Literaturrecherche zeigte ein kontinuierliches Anwachsen der Wolfspopulation in den letzten Jahren und insbesondere seit 2016 einen deutlichen Anstieg wolfsverursachter Nutztierschäden, wobei Pferde selten betroffen waren. Bei der Online-Befragung stammte die Hälfte der 574 ausgewerteten Fragebögen aus Brandenburg und Niedersachsen. Den größten Einfluss auf die Gefährdungsbeurteilung eigener Pferde durch die wachsende Wolfspopulation hatte das Wissen der Pferdehalter über Wolfsangriffe im eigenen Landkreis, wobei insbesondere die Aspekte einer Haltung von Jungpferden und Weidegang für die Pferde eine Rolle spielten. 64 % der Befragten gaben an, ihre Pferdehaltung trotz der zunehmenden Wolfspopulation nicht geändert zu haben. Nur 8 von 576 Pferdehaltern gaben amtlich bestätigte Wolfsübergriffe an und 30 hatten einen vermuteten Wolfsschaden amtlichen Stellen nicht gemeldet. Mehr als die Hälfte der Befragten, die Kontakt zu einem Wolfsberater hatten, bezeichneten die Zusammenarbeit als nicht bis wenig zielführend. Schlussfolgerung und klinische Relevanz Die Zahl amtlich bestätigter Wolfsangriffe auf Pferde ist gering. Durch einen grundsätzlich durchgeführten Gentest bei entsprechendem Verdacht ließen sich diese Zahlen objektivieren. Trotz des Bewusstseins einer zunehmenden Gefährdung von Pferden durch Wölfe unternehmen Pferdehalter überwiegend keine prophylaktischen Schutzmaßnahmen. Die Kommunikation zwischen den für das Wolfsmonitoring zuständigen Behörden und den Pferdehaltern erscheint verbesserungswürdig.
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TRY TO IMAGINE a small group of wolves sitting at a table engaged in vigorous debate. These wolves are from various parts of the globe and are perhaps a bit more scholarly than most. In fact, they are especially knowledgeable about the biology of that notorious two-legged species, Homo sapiens. They have been brought together to document their relationship with humans over the last several millennia. Pause for a few moments and consider what they might say ... Perhaps the wolves' discussion would chronicle the evils of the human species, including details of atrocities committed against lupine ancestors down through the centuries. They might discuss the bizarre workings of the human imagination and the hopeless confusion of fact and fiction about wolf relationships with humans. The discussion might also express admiration for the way early humans respected wolves and imitated their living in family bands, maintaining pair bonds for years at a time, communicating in complex ways, and hunting cooperatively. The effects of advances in human technology might be detailed. The recent and long-awaited legal protection for wolves and the soaring popularity of wolves among some humans would certainly deserve mention. After an exhaustive review of the wolf-human relationship, the wolves might finally conclude that it has taken so many forms, depending on time and place, that generalizations are impossible.
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Military training areas (MTA) are increasingly recognized as sites that harbor high levels of biodiversity, including large numbers of endangered species, yet their conservation value has not been rigorously assessed. Here, we studied the species richness and composition of medium- and large-sized mammals in Formosa MTA, a Brazilian military area, as a case study to assess the conservation value of military areas. We also made an evaluation of Brazilian MTAs regarding size, distributions, and representation of within ecoregion/biome compared with other protected areas. The medium- and large-sized mammal community composition fits the pattern described for the Cerrado, characterized by species of wide distributions, but locally rare. The Formosa MTA supports a relatively higher richness (n = 29) than protected areas in nearby regions and is a refuge for eight endangered species. Our study identified 52 MTAs covering a total area of 3 million ha. Our findings highlight the relevance of Formosa MTA for the conservation of regional mammalian fauna and indicate the potential of other military areas in the context of biological conservation.
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Roads fragment landscapes and trigger human colonization and degradation of ecosystems, to the detriment of biodiversity and ecosystem functions. The planet’s remaining large and ecologically important tracts of roadless areas sustain key refugia for biodiversity and provide globally relevant ecosystem services. Applying a 1-kilometer buffer to all roads, we present a global map of roadless areas and an assessment of their status, quality, and extent of coverage by protected areas. About 80% of Earth’s terrestrial surface remains roadless, but this area is fragmented into ~600,000 patches, more than half of which are <1 square kilometer and only 7% of which are larger than 100 square kilometers. Global protection of ecologically valuable roadless areas is inadequate. International recognition and protection of roadless areas is urgently needed to halt their continued loss. Read more on ...
Technical Report
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The present guideline for dealing with wolves (Canis lupus) in Germany was elaborated in the project “Scientific concept for a wolf management in Germany” initiated by the Federal Agency for Nature Conservation. It is supposed to be the scientific basis on which to develop a management plan for wolves in Germany, that has to be coordinated among and agreed on by the authorities and the interest groups. The first part of the manual contains background information concerning the status of wolves in Germany and Europe, as well as management approaches already applied. The second part presents in detail the scientific basis and recommendations for a wolf management in Germany. Currently reproducing wolves are restricted to North-Eastern Saxony, bordering Bran-denburg and Poland. The nearest larger wolf population is found in Eastern Poland. In the course of the project a human dimensions study was conducted, showing that the majority of Germans has a positive attitude towards wolves. However, a minority is strongly against wolves. Analysis of all articles published since 2001 in daily or weekly newspapers revealed, that the clear majority of articles had a neutral or positive tenor. The distribution of areas in Germany that resemble those that are inhabited by wolves today concerning habitat, overdevelopment and population density was analysed in a GIS study. Especially in the Northeast, the low mountain areas and the Southeast of Germany similar areas are widespread. Furthermore it was analysed for the whole country on district level which potential for conflicts has to be expected if wolves estab-lish themselves. In this respect the number of livestock and the way they are kept (if known) as well as the relative abundance of ungulates were compared for each district. The occurrence of wolves in the border area and even more the expected expansion of the strictly protected species, needs a coordinated approach of the states and nations involved. The goal envisaged by law is a viable German-Western Polish wolf population. The guideline recommends for Germany a pragmatic approach to reach this goal. Interference with the development of the population should be kept to a minimum. The wolf population should constantly be closely monitored by scientific standards. Conflicts that occur should be identified early on and minimized as much as possible. Of overall importance are the implementation of a nationwide coordinated concept for prevention and compensation of livestock damages, an intensive engagement in public relations and a close co-operation with the relevant interest groups. The manual analyses the complex and problematic issue of hunters, wolves and game species and recommends ways to minimize the conflicts. The question of public safety is thoroughly dealt with, the problem of possible hybridisation between wolves and dogs is discussed and recommendations are given for the handling of injured and dead wolves. Also aspects of keeping wolves and wolf-dog-hybrids in captivity are addressed. Last but not least, the guideline deals with the problem of illegal actions against wolves and stresses the need to push though the legal requirements. Concrete suggestions are given for the long-term organisation of monitoring and management of wolves in Germany, for international co-operation and applied research.
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Centuries of persecution have influenced the behaviour of large carnivores. For those populations persisting in human-dominated landscapes, complete spatial segregation from humans is not always possible, as they are in close contact with people even when they are resting. The selection of resting sites is expected to be critical for large carnivore persistence in human-dominated landscapes, where resting sites must offer protection to counteract exposure risk. Using wolves (Canis lupus) as a model species, we hypothesised that selection of resting sites by large carnivores in human-dominated landscapes will be not only influenced by human activities, but also strongly determined by cover providing concealment. We studied the fine-scale attributes of 546 wolf resting sites and confronted them to 571 random points in NW Iberia. Half of resting sites (50.8 %) were found in forests (mainly forest plantations, 73.1 %), 43.4 % in scrublands, and only 5.8 % in croplands. Compared to random points, wolves located their resting sites far away from paved and large unpaved roads and from settlements, whereas they significantly selected areas with high availability of horizontal (refuge) and canopy cover. The importance of refuge was remarkably high, with its independent contribution alone being more important than the contribution of all the variables related to human pressure (distances) pooled (51.1 vs 42.8 %, respectively). The strength of refuge selection allowed wolves even to rest relatively close to manmade structures, such as roads and settlements (sometimes less than 200 m). Maintaining high-quality refuge areas becomes an important element to favour the persistence of large carnivores in human-dominated landscapes as well as human-carnivore coexistence, which can easily be integrated in landscape planning.
Aim To compare predictions of the habitat suitability model (HSM) for wolves Canis lupus in Poland with actual wolf distribution in western Poland after 15 years of recolonization. Location Western Poland (WPL, ca. 136,000 km2), west of the 18°48′E meridian. Methods Data on wolf occurrence (8,057 records) were gathered in 2001–2016. Wolf presence in 10 × 10 km cells was classified as follows: (1) permanent occurrence with reproduction, (2) permanent occurrence with no reproduction and (3) sporadic occurrence (interpreted as dispersing individuals). These cells were compared to all 10 × 10 km cells in WPL with respect to the probability of wolf occurrence as predicted by the HSM and habitat variables important for wolves. For temporal analysis, data were divided into two 8-year subsets: the initial and later phases of wolf recovery. Results Wolves were recorded in 259 cells (19.8% of the study area). The pairs and packs settled in areas predicted by the HSM to have good and very good habitat, in cells characterized by high forest cover and low densities of roads. Wolf groups that reproduced were found in the best-quality habitats characterized by denser forest cover and markedly lower shares of anthropogenic structures. Dispersing individuals were mostly recorded in unsuitable and suboptimal habitats, and they avoided both the poorest and the best habitats. In the initial phase of wolf recovery, cells selected by wolves for settling down and those used by dispersing wolves did not differ in their habitat parameters. However, in the later phase, as WPL became more saturated with wolf packs, dispersing individuals were recorded in less suitable habitats. Main conclusions The HSM for Polish wolves predicted with high accuracy the areas later occupied by wolf groups in the western part of the country. A similar approach may also be useful to predict the future distribution of wolves in the lowlands of central and western Europe where environmental conditions are comparable and recolonizing wolves originate from the same source population.
Military training areas (MTAs) cover 6% of the earth's land surface, but the impact on biodiversity of weapons use in MTAs remains largely unknown. We quantified the effects of military training on vertebrates in a 5-year study at Beecroft Weapons Range in south-eastern Australia by contrasting the occurrence of birds, mammals and reptiles between 24 sites within an area subject to repeated weapons use and a matched set of non-impacted sites. Species richness of mammals and reptiles was similar within versus outside the impact area, although many individual species responded to fire, which occurred more frequently in impacted sites. Bird species richness, the occurrence of larger-bodied and migratory bird species, and the occurrence of most individual bird species, was reduced within the impact area. Many bird species that displayed low prevalence in impacted sites also declined over time across the whole study area. Differences in biota between the impact and non-impact areas were detectable after controlling for the effects of recent fire, suggesting that weapons use impacted vertebrates through mechanisms additional to altered fire regimes. Overall, our data indicated that Beecroft Weapons Range maintained considerable biodiversity value despite prolonged military use. Hence, MTAs have the potential to make a substantial contribution to conservation outside the formal protected area network. However, managers of MTAs need to explicitly state their environmental objectives. This is because management practices may be different if the aim is to maximize species richness rather than to secure populations of particular species.