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Received: 20 July 2018 Revised: 12 October 2018 Accepted: 27 January 2019
DOI: 10.1111/conl.12635
LETTER
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,
Germany
2Goethe-University Frankfurt am Main,
Department of Biological Sciences, Germany
3Senckenberg Research Institute and Natural
History Museum Frankfurt, Gelnhausen,
Germany
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,
Germany
Correspondence
Ilka Reinhardt, LUPUS – German Institute
for Wolf Monitoring and Research, Dorfaue 9,
D-02979 Spreewitz, Germany.
Email: ilka.reinhardt@wolves-germany.de
Funding information
Robert Bosch Stiftung; Regina Bauer Stiftung
Abstract
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.
KEYWORDS
Canis lupus, large carnivores, population growth, protected areas, recolonization
1INTRODUCTION
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. wileyonlinelibrary.com/journal/conl 1of7
https://doi.org/10.1111/conl.12635
2of7 REINHARDT ET AL.
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.
2METHODS
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
REINHARDT ET AL.3of7
(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,
2017).
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.
3RESULTS
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
4of7 REINHARDT ET AL.
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).
4DISCUSSION
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
pack.
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
REINHARDT ET AL.5of7
TABLE 1 Number of territory years (cumulative number of years territories were occupied) and the number of wolves found dead within
territories
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
6of7 REINHARDT ET AL.
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.
ACKNOWLEDGEMENTS
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.
AUTHOR CONTRIBUTIONS
IR and TM conceptualized the paper, IR analyzed the data,
and all authors contributed to the writing of the manuscript.
CONFLICT OF INTERESTS
The authors declare no conflict of interests.
ORCID
Thomas Mueller https://orcid.org/0000-0001-9305-7716
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SUPPORTING INFORMATION
Additional supporting information may be found online in the
Supporting Information section at the end of the article.
How to cite this article: Reinhardt I, Kluth G, Nowak
C, et al. Military training areas facilitate the recolo-
nization of wolves in Germany. Conservation Letters.
2019;e12635. https://doi.org/10.1111/conl.12635
