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Conflicts with Wolves Can Originate from Their Parent Packs

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Transmission of experience about prey and habitat supports the survival of next generation of wolves. Thus, the parent pack (PP) can affect whether young migrating wolves (loners) kill farm animals or choose to be in human environments, which generates human–wolf conflicts. Therefore, we researched whether the behavior of loners resembles PP behavior. After being extinct, 22 loners had entered the Netherlands between 2015 and 2019. Among them, 14 could be DNA-identified and linked with their PPs in Germany. Some loners were siblings. We assessed the behavior of each individual and PP through a structured Google search. PP behavior was determined for the loner’s rearing period. Similarity between loner and PP behavior was significant (p = 0.022) and applied to 10 of 14 cases: like their PPs, three loners killed sheep and were near humans, five killed sheep and did not approach humans, while two loners were unproblematic, they did not kill sheep, nor were they near humans. Siblings behaved similarly. Thus, sheep killing and proximity to humans may develop during early-life experiences in the PP. However, by negative reinforcement that can be prevented. New methods are suggested to achieve that. As a result, new generations may not be problematic when leaving PPs.
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Animals 2021, 11, 1801.
Conflicts with Wolves Can Originate from Their Parent Packs
Diederik van Liere 1,*, Nataša Siard 2, Pim Martens 3 and Dušanka Jordan 2
1 Institute for Coexistence with Wildlife, Heuvelweg 7, 7218 BD Almen, The Netherlands
2 Department of Animal Science, Biotechnical Faculty, University of Ljubljana, Groblje 3,
1230 Domžale, Slovenia; (N.S.); (D.J.)
3 Maastricht Sustainability Institute, Maastricht University, P.O. Box 616, 6200 MD Maastricht,
The Netherlands;
* Correspondence:
Simple Summary: Conflicts with wolves arise because wolves kill farm animals, especially sheep,
or approach humans. It is expected that young wolves learn from their parent pack (PP) what their
prey is and if it is safe to be near humans. To confirm this, we researched whether the behavior of
young migrating wolves (loners), after they leave the pack, resembles PP behavior. Fourteen loners
entering the Netherlands between 2015 and 2019 could be identified and genetically linked to their
PPs. Loner and PP behavior was similar in 10 out of 14 cases. Like their PPs, some young wolves
killed sheep and were near humans, others killed sheep and did not approach humans, while two
loners were unproblematic, they did not kill sheep nor were they in proximity to humans. Thus, the
PP behavior did predict loner’s behavior and conflicts may be similar between young wolves and
their PPs. However, conflicts need not arise. To achieve that, new prevention methods are pro-
posed to teach wolves in the PP not to approach sheep and humans. As a result, new generations
may not be problematic when leaving the PP.
Abstract: Transmission of experience about prey and habitat supports the survival of next genera-
tion of wolves. Thus, the parent pack (PP) can affect whether young migrating wolves (loners) kill
farm animals or choose to be in human environments, which generates human–wolf conflicts.
Therefore, we researched whether the behavior of loners resembles PP behavior. After being ex-
tinct, 22 loners had entered the Netherlands between 2015 and 2019. Among them, 14 could be
DNA-identified and linked with their PPs in Germany. Some loners were siblings. We assessed the
behavior of each individual and PP through a structured Google search. PP behavior was deter-
mined for the loner’s rearing period. Similarity between loner and PP behavior was significant (p =
0.022) and applied to 10 of 14 cases: like their PPs, three loners killed sheep and were near humans,
five killed sheep and did not approach humans, while two loners were unproblematic, they did not
kill sheep, nor were they near humans. Siblings behaved similarly. Thus, sheep killing and prox-
imity to humans may develop during early-life experiences in the PP. However, by negative rein-
forcement that can be prevented. New methods are suggested to achieve that. As a result, new
generations may not be problematic when leaving PPs.
Keywords: human–animal conflict; wolf behavior; migrating wolves; sheep killing; early-life
experiences; bold wolves; learning; depredation; deterrence
1. Introduction
Wolves in Europe live in territorial packs of about 2–7 animals. A pack resembles a
family and usually consists of a breeding pair, their offspring from previous years, and
sometimes unrelated wolves. All members of the family pack cooperate in raising the
cubs, i.e., they protect them, feed them [1], and train them in social, hunting, and survival
skills [2]. Young wolves remain in the pack for at least the first 10 months, but usually
disperse between 1 and 2 years of age [3] to find mates or a territory of their own [1,4].
Citation: van Liere, D.; Siard, N.;
Martens, P.; Jordan, D. Conflicts
with Wolves Can Originate from
Their Parent Packs. Animals 2021, 11,
Academic Editor: Pia Lucidi
Received: 21 April 2021
Accepted: 13 June 2021
Published: 16 June 2021
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Animals 2021, 11, 1801 2 of 20
They are then referred to as loners. Compared to other social canids, wolves are parented
for a relatively long time [5]. The cubs suckle milk and feed on partially digested food
regurgitated by the parents or elder siblings until weaning at 10 weeks of age. During
these weeks, they receive more and more carcass pieces and complete, but small, opened
carcasses [6]. In this way, they become accustomed to the smell and taste of the prey that
constitutes their diet [7]. Food can be not only parts of dead animals, but also plant matter
such as berries and mushrooms [8]. Young wolves tend to imitate pack members [9] and
around the time of weaning, they begin to follow pack members to carcasses [4,6]. Young
wolves observe how prey is killed, and as they become stronger, they actively participate
and train to approach and kill prey [7,10]. Indeed, Radinger [11] observed that early in
life, wolves learn from their parents and elder siblings what kind of prey to select and
kill. Thus, young wolves develop a keen knowledge of what prey is, as well as habits and
search patterns that increase their hunting efficiency [1,12,13] and their chance of survival
[14]. The development of prey perception, selection, and its coupling with the develop-
ment of prey catching skills applies to mammalian predators in general [15,16]. It is also
generally accepted that the transmission of experience about prey and habitat is instru-
mental in preparing a new generation. Consequently, the new generation prefers a hab-
itat comparable to the one in which it was reared. Thus, the vertical transmission of ex-
perience enables the new generation to survive efficiently [4,17–20].
Wolves living near people are described as cautious about where and when to travel
[21] and tend to retreat when they detect approaching humans [22]. However, under
certain circumstances, wolves can lose their fear of humans [23,24]. To prevent aggres-
sion towards humans, it is important to keep wolves in the wild and maintain their fear
[23]. According to Karlsson et al. [22], wildlife managers should detect wolves that be-
have boldly toward humans during the early stages of habituation. However, they state
that the development of boldness is poorly studied. Dispersing wolves would prefer a
habitat, depending on early life experiences [25,26], as is known for coyotes [27]. There-
fore, a young wolf’s preference for human habitat may also have evolved during the
rearing period, especially if the pack is not fearful of humans. In addition, attraction to
humans or human habitat can be based on positive reinforcement by food, dogs, or play
opportunities [22,24].
The last Dutch wolf was killed in 1845 [28], but after this extinction, since March
2015, numbers of loners have entered the Netherlands. The habitat they have entered
consists of densely populated delta lowland including a dense transport infrastructure,
intensive agriculture and urban expansion, and fragmented nature areas. It covers 41.5
thousand km2: built-up area and roads cover 15%, forests and open natural areas cover
12%, 19% is water, and 54% is agricultural land [29]. Potential wild prey such as roe deer
is present almost everywhere in the Netherlands, but with lower distribution in the
coastal provinces. In 2015 the number of roe deer in the Netherlands would be up to 100
thousand [30]. As agricultural areas (excluding glasshouse area) and natural areas (ex-
cluding waters) sum to 27.2 thousand km2 [29], the density is estimated to be 3.7 roe deer
per km2. The other Dutch wild ungulates (red deer, fallow deer, and wild boar) only live
in restricted, fenced areas, the largest being the Veluwe (about 1000 km2).
The recent observations of loners have been along roads, in agricultural fields and in
villages, or identified when found as road kills, or from DNA samples, e.g., from kills of
farm animals. Identification of individual wolves is performed in a current wolf DNA
sampling program by the Central European wolf (CEwolf) consortium [31], a coalition of
Dutch, Belgian, and German scientists, amongst others. This program also makes it pos-
sible to identify their parent packs. Therefore, there is a unique opportunity to test the
hypothesis that habits in prey and habitat choice are similar between parent packs and
loners. That is the aim of this paper. If there is a high degree of similarity, this study also
has practical implications. Sheep were the only livestock species attacked in the Nether-
lands in the period 2015–2019 [32] and (together with goats) the most preyed livestock
species by wolves across Europe [33]. Therefore, whether a parent pack kills sheep may
Animals 2021, 11, 1801 3 of 20
predict what a loner would prefer and predict whether the loner would become prob-
lematic towards sheep. This prediction differs from the common assumption that sheep
are easy prey [8,34–36]. Similarly, whether loners will be in proximity to a human envi-
ronment may also relate to a habit of their parent packs.
2. Materials and Methods
Twenty-two wolves had entered the Netherlands as loners between March 2015 and
March 2019. Of these, 17 were identified by DNA analysis and these are the subject of our
study. As wolves are new in the Netherlands, the sight of a wolf, or the killing of farm
animals was closely followed by the media, dedicated websites, and general public. For
DNA analysis, samples of feces or hair were collected from a site where a live wolf was
seen, or could be found, without observation of the actual wolf. Slime was collected from
wounds of killed farm animals. Tissue from dead wolves was also sampled. These sam-
ples had to be collected soon after the source was discovered because DNA quality can
decline rapidly under field conditions and cannot be reliably analyzed after two days
[37]. DNA analysis of these samples was performed and published by the teams of An-
imal Ecology of Wageningen University, the Netherlands, Research Institute for Nature
and Forest, Geraardsbergen, Belgium and Naturschutzgenetik of the Senckenberg For-
schungsinstitut, Gelnhausen, Germany. These teams have standardized their DNA
analysis [38] and are collaborating in the CEwolf consortium. They identify wolves, i.e.,
assign GW (grey wolf) numbers, and register individual wolves and reproducing packs.
Genotyping is described by Harms et al. [37] and the Senckenberg Forschungsinstitut
[39]. Control-region sequences of mitochondrial DNA are analyzed to verify if it is a wolf
species. If so, identification of each individual is possible by additional microsatellite and
single nucleotide polymorphism (SNP) genotyping of biparentally inherited and patri-
lineal Y-chromosome markers [38,40]. The success rate of SNP genotyping is 87% [41].
The behavior of each identified loner was classified as (1) known to kill sheep or not,
within 6 months after the loner was detected in the Netherlands, and (2) known to be in
proximity to humans or not, also within 6 months after the loner was detected. We chose
the 6-month period, because we were interested in the choices of young wolves that are
relatively inexperienced, not territorial, and not yet developing their own loca-
tion-specific routines, and while migrating, probably mainly relying on experiences
gained in the parent pack. If the loner was involved in a farm animal attack this was ta-
bled in freely available Excel files on the website of the Dutch interprovincial BIJ12 de-
partment [32]. Kills of Dutch farm animals are always reported to the BIJ12 department,
as this department financially compensates the losses if wolf DNA was found in wound
swabs. The class “being in proximity to humans” was applicable whenever there were
eyewitness descriptions, photos, or videos indicating that a wolf had entered a human
settlement (village, town, etc.), or eyewitness descriptions, photos, or videos indicating
that a wolf had approached humans to a distance of 30 m or less, despite obvious op-
portunities to stay at a greater distance. The 30 m distance is defined by the Large Car-
nivore Initiative for Europe LCIE [24] as a close encounter between humans and wolves.
In relation to proximity, loner identities were determined based on the Wageningen
Animal Ecology team’s identification as reported in news publications. If a loner was
only sighted, but its DNA identity was not provided in the reference, we assumed it was
the DNA-identified wolf whose DNA track was found in or within 15 km of the sighting.
This was possible because a trail of subsequent nearby wolf sightings appeared in the
media suggesting that it was the same individual. The trail could match a repeated DNA
sample that verified the individual in question. The identity of a sighted wolf could not
be estimated if no DNA evidence was collected within 15 km of the sighting. We chose
the 15 km distance, as loners may pause migration and temporarily stay in a certain area
[42]. We considered this to be similar to a stay in a territory. The 15 km distance covers
the distances between the start and end sites of a travel day of Polish wolves in their ter-
Animals 2021, 11, 1801 4 of 20
ritory (max. 11 km [12]), as well as the maximum total daily distance between six site
measurements per day of German wolves in their territory (max. 13 km [42]).
Parent pack recognition was possible by microsatellite and SNP genotyping allow-
ing the detection of alleles stemming from both the mother and the father [37,39,43]. This
genotyping could match with the CEwolf consortium DNA profile database of known
wolf individuals and packs and enabled the consortium to trace parent packs of the
Dutch loners. Where known, parent packs of loners involved in farm animal attacks were
listed in the freely available BIJ12 Excel file [32]. We used the one published 4 January
2020. For loners not associated with farm animal kills, parent packs were disclosed by the
Wageningen Animal Ecology team in their publications on loners.
To find descriptions of the parent pack’s behavior in relation to sheep kill and
proximity to humans during rearing of the loners, the birth year of the loner was esti-
mated from autopsy reports [44–46], reports of tagged individuals caught while still with
the parents [47,48], or when the individual was first detected within the pack through
DNA analysis [49,50]. These data were combined with the first year of reproduction of
the parent pack [49], as well as the assumption that May is the month of birth [12]. If a
first DNA detection of a loner was revealed while still in the parent pack that had been
reproducing for several years and was frequently monitored by DNA sampling, it was
determined that this loner was born in May prior to the detection.
Descriptions of the selection of sheep as prey or proximity to humans by the parent
pack of the loners were found via Google search. Selection of sheep as prey was also
checked in government databases of farm animal kills [51–53]. A 10-month period was
chosen for descriptions of parent packs: between 1 May of the loner’s birth year and 1
March of the following year. Births take place in May [12], so the mentioned period co-
vers the usual minimum period during which young wolves remain in the pack [3].
During this period the loner is raised and gets its first experiences regarding prey and
possible proximity to humans by the pack [4]. Since all loners that entered the Nether-
lands stemmed from German packs, we used the German words for pack (Rudel), farm
animal damage (Nutztierschäden), farm animal attack (Nutztierrisse), sheep killed
(Schafe getötet), near people (Nähe Menschen), and not being shy (keine Scheu) as key-
words. Thus, the following search strings were used: “Rudel” and “location parents” and
“year of birth” (for example “Rudel Ueckermünder Heide 2014”). We also used “Rudel”
and “location parents” and “Nutztierschäden” and “year of birth”, “Rudel” and “loca-
tion parents” and “Nutztierrisse” and“year of birth”, “Rudel” and “location parents” and
“Schafe getötet” and “year of birth”, “Rudel” and “location parents”, and “Nähe
Menschen” and “year of birth”, “Rudel” and “location parents” and “keine Scheu” and
“year of birth”. We additionally combined these keyword sequences with the
GW-number of the involved loner or the GW-numbers of the respective parents. All of
the first 20 hits were read and filtered with respect to evidence of sheep killing, entering
human settlements, or being within 30 m of humans. If a filtered hit contained the loca-
tion of a sheep kill or a wolf observation, the parent pack involved was classified as
“known to kill sheep” or “known to be in proximity to humans”. The pack was identified
either by DNA or by the distance between the location of the sheep kill or observation,
and the center of that pack’s territory, if DNA identification was not revealed in the ref-
erences. The location of pack territory was provided by the Dokumentations- und
Beratungsstelle des Bundes zum Thema Wolf DBBW [54] in a map with territory loca-
tions as circles with a scaled radius of 8 km. Although territory size varies between and
within packs depending on season and prey abundance [12,42], the area of repeated
presence of German pack members was determined by DNA analyses and/or repeated
independent sightings by wildlife experts [55]. The German authorities then translated
these results into the aforementioned published circles as indications of where the packs
were located. The center of this circle was used to measure the distance between the
parent pack and the location of a kill or observation. A range of 15 km was chosen for the
same reasons as described above for the loners. Within this range, it was assumed that
Animals 2021, 11, 1801 5 of 20
the kill or observation involved wolves from the pack. If there was more than one pack
near the kill or observation site, it was assumed that the kill or observation involved
wolves from the nearest pack. If Google searches on a parent pack within the period of
May–March, which begins in the loner’s birth year, did not reveal any evidence of sheep
killing or proximity to humans, the parent pack was classified as “not known to kill
sheep” or “not known to be in proximity to humans”, respectively. For some loners the
birth year could remain uncertain and could be one or two years before the first discov-
ery in the Netherlands. If this was the case, we looked for descriptions of prey choice and
proximity by members of the parent pack for the period between 1 May and 1 March in
both possible birth years. During these periods, any description of sheep kill or proximity
that met the above criteria was used to classify the parent packs of these loners. Only
when no descriptions were found were they classified as “not known to kill sheep” or
“not known to be in proximity to humans”.
For the parent pack and the loners four classifications could apply: (1) no kill of
sheep, no proximity to humans, (2) no kill of sheep, proximity to humans, (3) kill of
sheep, no proximity to humans, and (4) kill of sheep, proximity to humans. The classifi-
cation of the parent pack as independent variable and that of the loner as dependent
variable was analyzed in a multinomial logistic regression. In addition, both classifica-
tions were cross-tabulated and Cramér’s V coefficient for their association [56] was cal-
culated. IBM-SPSS statistics version 25 was used for these calculations.
3. Results
Between March 2015 and March 2019, 17 non-settled loners entering the Nether-
lands were detected and identified (Table 1). The median interval of entry of successive
loners into the Netherlands was 1.5 months (min: 0 and max: 18; Table 1). Only once were
two different loners detected in the Netherlands for the first time on the same day. The
distance between the locations of these first detections was 62 km. For 14 of these loners,
a parent pack could also be traced back, which was always in Germany. There were 11
parent packs involved: seven were in Niedersachsen, two in Mecklenburg-Vorpommern,
one in Sachsen, and one in Brandenburg (Table 1). Three pairs of loners were related: two
females stemmed from the Ueckermünder Heide (nr 2 and 17); nr 2 was observed in
September 2016, nr 17 in March 2019. Two males were from the Barnstorfer Moor pack
(nr 6 and 7). They were observed in February and March 2018. One male and one female
were from Babben-Wanninchen (nr 4 and 11). The male was first observed in October
2017 and the female in May 2018. The year of birth could be reliably estimated for 11
loners (nr 1, 3, 4, 5, 6, 7, 11, 13, 15, 16, 17). The median age of these loners was 20 months
(min: 9; max: 30; Table 1).
Animals 2021, 11, 1801 6 of 20
Table 1. Identified loners that have entered the Netherlands (NL), numbered chronologically with their ID number (nr)
based on DNA analyses (f: female; m: male), references of loners including ID numbers, date and location of first detec-
tion in the Netherlands, age, year of birth and method of age determination (G: first genetic detection; A: autopsy; C:
capture in the pack; F: first litter; without a letter: uncertain) and location of parent pack (N: Niedersachsen; MV: Meck-
lenburg-Vorpommern; B: Brandenburg; S: Sachsen; n.k.: not known).
Loner Nr
with ID Nr
Date First
Detection NL
Location First
Observation NL
Age in Months/
Year of Birth Parent Pack/Land
in Germany
1 GW386m [32,57] 07/03/15 Bargerveen 10/2014G Truppenübungsplatz in
Munster Nord/N
2 a GW620f [49,58] 03/09/16 Beuningen >16/2014 or 2015 Ueckermünder Heide/MV
3 GW657m [46] 03/03/17 Veeningen near
highway A28 22/2015A Cuxhaven Langes Moor/N
4 b GW843m [32,45] 14/10/17 Laag Zuthem 30/2015A Babben Wanninchen/B
5 GW680f [32,47,59,60] 19/12/17 Forest Hardenberg 18/2016C Lübtheener Heide/MV
6 c GW955m [32] 21/02/18 Betuwe area, river
Nederrijn 9/2017F Barnstorfer Moor/N
7 c GW913m [32,44] 05/03/18 Lottum- Venlo area 10/2017A Barnstorfer Moor/N
8 GW954f [32] 18/03/18 Benneveld >10/2016 or 2017 Schneverdingen/N
9 GW953m [32] 18/03/18 Beckum n.k. n.k.
10 GW763f [32] 04/04/18 Boijl >11/2015 or 2016 Daubitz/S
11 b GW998f [32,61,62] 03/05/18 Langezwaag 24/2016C Babben Wanninchen/B
12 GW979m [32,63,64] 16/06/18 Buitenpost n.k. n.k.
13 GW960f [62] 01/08/18 Midden Veluwe 14/2017G Göhrde pack/N
14 GWxxxf [32,62] 03/10/18 Orvelte n.k. n.k.
15 GW893m [32,65] 05/01/19 Damsholte 20/2017G Eschede/Rheinmetall/N
16 GW965f [32,66] 22/02/19 Lemelerveld 21/2017G Die Lucie/N
17 a GW849f [32,48] 29/03/19 Hooghalen 21/2017C Ueckermünder Heide/MV
a, b, c: same letters are siblings.
Fourteen loners were responsible for attacks on sheep between March 2015 and
March 2019 (Table 2); 203 kills were recorded [32]. Table 2 summarizes references that
describe the loners killing sheep and proximity to humans, or not.
Table 2. Identified loners and references with evidence of killing sheep and proximity to humans
within 6 months after the first detection in the Netherlands.
Loner Nr Loner’s ID Nr Killing of Sheep Proximity
1 GW386m Yes [32] Yes [50,67–69]
2 a GW620f No No
3 GW657m No No
4 b GW843m Yes [32,45] No
5 GW680f Yes [32,70] No
6 c GW955m Yes [32] Yes [71–73]
7 c GW913m Yes [32,74] Yes [74,75]
8 GW954f Yes [32] No
9 GW953m Yes [32] No
10 GW763f Yes [32] No
11 b GW998f Yes [32,61,62] No
12 GW979m Yes [32,63] No
13 GW960f No No
14 GWxxxf Yes [32,76] No
15 GW893m Yes [32,65] No
16 GW965f Yes [32,66] No
17 a GW849f Yes [32] No
a, b, c: same letters are siblings.
Animals 2021, 11, 1801 7 of 20
Three of the 17 loners (18%) were not associated with killing sheep in a 6-month pe-
riod following their detection (wolf nr 2, 3, and 13, Table 2). Wolf nr 2 was photographed
from a distance in September 2016 defecating in an open farmland at daylight [77]. DNA
was taken from feces collected at that spot [49,58]. DBBW [49] reported that this female
was from the German pack Ueckermünder Heide in Mecklenburg-Vorpommern. She had
also been detected in Engden, Germany, in August 2016 [78]. After the Dutch encounter,
she was never detected again. Wolf nr 3 was killed by a car in March 2017. His DNA was
also never linked to the killing of farm animals. The autopsy revealed that his stomach
contained remains of hare [46]. Wolf nr 13 was only detected by feces samples collected
in August 2018 in the Midden Veluwe nature area. It turned out that she settled there, as
her DNA was repeatedly found thereafter in this area for 6 months in a row.
Three identified loners (nr 1, siblings 6 and 7) were observed in proximity to humans
(near people and walking through villages). They also killed sheep (Table 2) and formed
together 21% of all identified wolves that killed sheep. Wolf nr 1 was recorded traveling
through Niedersachsen in Germany and near people since February 2015 [50] and en-
tered the Netherlands in March 2015. He was filmed walking along the footpath of the
main road through the Dutch village of Kolham [67] (Figure 1), on several occasions
walking alongside cars, crossing fields and roads near people and cars in broad daylight.
Figure 1. Satellite image overviewing the village Kolham and the trajectory (in red) from south to north of wolf 1 when
filmed. Image: ©Google Earth.
Wolf nr 6 was recorded walking through the center of several villages such as in
Bennekom where on 21 February 2018 this wolf was filmed twice [71] and subsequently
filmed the same night in Veenendaal [79] at 9 km distance (Figure 2), passing cyclists and
cars. This wolf was seen both in broad daylight and at night.
Animals 2021, 11, 1801 8 of 20
Animals 2021, 11, 1801 9 of 20
Figure 2. Satellite image overview (a) of the subsequent east to west video-recorded locations of wolf 6 in one night: (b) in
red and pink in Bennekom; (c) in blue in Veenendaal. Images: ©Google Earth.
His brother, wolf nr 7, was repeatedly seen in Belgian villages Dilsen-Stokkem and
Meeswijk [80,81]. All other loners were never reported to be seen in proximity to hu-
The behavior of the parent pack in relation to killing sheep and proximity to humans
at the time that a loner was born is described in the references in Table 3. No references of
sheep killing were found for two parent packs (relating to loner nr 10 and 13). For the
other nine parent packs there are references of sheep killing. DNA proof on killed sheep
applied to five of these nine (relating to loner nr 3, brothers 6 and 7, and loners 8, 15, 16).
For the remaining four parent packs (relating to loner nr 1, sisters 2 and 17, siblings 4 and
11, and loner nr 5), references related to the distance between the site of a sheep kill and
the center of the parent pack territory were used. However, the sheep killing results of
two of these parent packs were not consistent in the course of time. This applied to the
parent pack Ueckermünder Heide (relating to loner nr 2 and 17) and the parent pack
Babben Wanninchen (relating to loner nr 4 and 11). The parent pack Ueckermünder
Heide started to reproduce in 2014; loner nr 2 was born either in 2014 or 2015 (Table 1).
No sheep kills could be associated with this pack in these years (Table 3), whereas in
2017, the birth year of loner nr 17 (Table 1), sheep kills occurred within 15 km of the ter-
ritorial center of this pack [82,83]. The parent pack Babben Wanninchen started to re-
produce in 2013. Loner nr 4 was born in 2015. The only kill of sheep during his rearing
period from May 2015 to March 2016 occurred in January 2016 [84], but this was 5 km
from the center of a neighboring pack and 13 km from the Babben Wanninchen parent
pack. Therefore, this kill was not attributed to the Babben Wanninchen pack (Table 3). As
of 2016, there was another father. Loner nr 11 was born in 2016 (Table 1). In October 2016,
there were two attacks on sheep within 6 km of the center of the nearest pack, namely the
Babben Wanninchen pack [84]. Thus, here we attributed sheep kill to this parent pack
(Table 3).
Animals 2021, 11, 1801 10 of 20
Table 3. Fourteen identified loners that were genetically linked to a parent pack, with their ID number, and references to
their parent pack regarding location (N: Niedersachsen; MV: Mecklenburg-Vorpommern; B: Brandenburg; S: Sachsen),
killing of sheep and proximity to humans in the period May of the loner’s birth year (or possible birth years) to March of
the next year.
Loner Nr Loner’s ID Nr Location/Land in Germany Killing of Sheep Proximity
1 GW386m Truppenübungsplatz, Munster/N Yes [52,85] Yes [50,69,86]
2 a GW620f Ueckermünder Heide/MV No No
3 GW657m Cuxhaven Langes Moor/N Yes * [52] No
4 b GW843m Babben Wanninchen/B No No
5 GW680f Lübtheener Heide/MV Yes [87,88] No
6 c GW955m Barnstorfer Moor/N Yes * [52,89] Yes [90]
7 c GW913m Barnstorfer Moor/N Yes * [52,89] Yes [90]
8 GW954f Schneverdingen/N Yes * [52] Yes [91]
10 GW763f Daubitz/S No No
11 b GW998f Babben Wanninchen/B Yes [84] No
13 GW960f Göhrde pack/N No No
15 GW893m Eschede/Rheinmetall/N Yes * [52] No
16 GW965f Die Lucie/N Yes * [52] No
17 a GW849f Ueckermünder Heide/MV Yes [82,83] No
*: DNA analysis demonstrated parent pack was involved in killing sheep; a, b, c: same letters are siblings.
The parent pack of loner nr 1, born in 2014 at the Truppenübungsplatz, Munster,
Niedersachsen (Table 1), was actually not related to references describing the killing of
farmed sheep in 2014/2015. Nevertheless, we assumed an involvement of the pack in the
killing of sheep (Table 3), because the pack is located in the Lünerburger Heide. This
heather habitat is well known for the free-ranging herds of a sheep breed called
Heidschnucke [92]. It is likely that the parent pack has experienced these sheep. Actual
wolf kills of such sheep in this area must have been occurring, as this motivated shep-
herds to install night enclosures in 2015. A lack of reference in the database of farm kills
in Niedersachsen can be explained as reporting and DNA sampling of killed free-ranging
Heidschnucke was not considered useful and was not performed [93]. An additional
argument is that the pack was known to kill sheep in previous years [85].
References of proximity to humans were found for three of the 11 parent packs (of
loner nr 1, brothers 6 and 7, and loner nr 8). These involved sites of proximity within 15
km of the center of the parent pack’s territory. In the case of loner nr 1, his parents and
siblings of Truppenübungsplatz, Munster pack were well known for their proximity to
humans [50,69]. Moreover, a sibling born in 2015, that remained in Germany,
ID-numbered as GW369m, and was not one of the Dutch loners, was fitted with a
VHF-transmitter around his neck. This wolf also repeatedly followed humans and at-
tacked dogs that were being walked. For this reason, he was shot [50]. In the second case,
the Barnstorfer Moor parent pack of sibling loners nr 6 and 7, eyewitness reports mention
wolves from this pack being in proximity [90]. In the third case, the Schneverdingen
parent pack of loner nr 8, a video from February 2016 shows a wolf approaching a wait-
ing car. This happened at a distance of 3 km from the center of their territory [91].
The characteristics of the parent packs at the time the loners were reared, and the
characteristics of the loners themselves have been summarized and cross-tabulated in
Table 4. These wolves only classified in the categories (1) no kill of sheep, no proximity to
humans, (3) kill, no proximity, and (4) kill and proximity. No parent pack or loner was
characterized by the class (2) no kill, proximity. The multinomial logistic regression
model was significant (p = 0.02) with behavior category of the parent packs as predictor
variable. Moreover, the Cramér’s V coefficient was 0.64, showing a significant association
between behavioral characteristics of the parent pack and of their loner (p = 0.02).
Animals 2021, 11, 1801 11 of 20
Table 4. Frequency distribution of sheep kill and proximity to humans characteristics of 14 identi-
fied Dutch loners and their parent packs at the time the loners were reared (bold: similar behavior;
loner nr in parentheses).
no sheep kill; no
proximity sheep kill; no proximity
sheep kill;
no sheep kill;
no proximity 2 (2, 13) 2 (4, 10)
sheep kill;
no proximity 1 (3) 5 (5, 11, 15, 16, 17)
sheep kill;
1 (8) 3 (1, 6, 7)
4. Discussion
Though based on a limited sample size, our results are in an agreement with the
hypothesis that there is a high degree of similarity in prey and habitat choice between
parent packs and their loners. They are similar regarding sheep killing or not and being
in proximity to humans or not.
DNA recognition allows reliable documentation of individual loners. However, not
all observations in our study are backed up by DNA identification, which could raise
doubt whether some data are correct. For example, for wolf 6, DNA samples were not
taken at each confrontation. Therefore, it can be questioned whether it was always the
same individual. Nevertheless, it is very unlikely that different individuals were present
at the same time and place on the subsequent days of the observations. First, there were
no resident wolves in the Netherlands until February 2019 [94]. Second, it is unlikely that
two loners migrated on the same track and at the same time, as the median time interval
between successive loners entering the Netherlands was 1.5 months. Even in the single
case of a first detection on the same day, the distance between loners was 62 km (loner 8
and 9, Table 1).
Another inaccuracy could be seen in the assessment of the parent pack involvement
when DNA identification is lacking and the distance between the site of a sheep kill or
proximity encounter and the center of the parent pack´s territory is used instead. The
shape and size of the territory at the time of the kill or proximity encounter was un-
known, the 15 km limit may not cover the actual exploration range of pack members, and
members from neighboring pack territories may explore the parent pack area [42]. Thus,
the possibility remains that a wolf involved was other than a parent pack one. Never-
theless, Google searches were a valuable tool to classify sheep kills or proximity to hu-
mans. We give some examples to illustrate this. For example, Koerner [69] and other
sources [50] describe that members of the parent pack of loner 1 Truppenübungsplatz in
Munster Nord have been in close proximity to humans repeatedly since 2012 and also
early 2015. Another example provided by Proplanta [82] and confirmed by Welt [83] de-
scribed the Ueckermünde as a regional hotspot for the killing of farm animals, including
sheep. This would apply to 2017 and 2018. This hotspot is 5 km from the center of the
parent pack Ueckermünde Heide, while loner nr 17 is born in 2017. Such reports of local
experiences contribute to the reliability of the classification of the parent pack, despite the
lack of DNA proof.
The repeated wolf reporting shows that the presence of wolves is highly profiled
and covered in the media in both Germany and the Netherlands, especially when wolves
are near humans or kill farm animals. This is partly due to unfamiliarity with wolves and
concern for the safety of people in a region with a new large predator. Wolves have been
reproducing in Germany only since 2000 [95,96], while the first proven Dutch wolf
presence was in 2015 (wolf nr 1), after more than 170 years of absence [28]. Despite po-
tential errors, the nominal measurements accumulated to a result in agreement with the
Animals 2021, 11, 1801 12 of 20
hypothesis. Improved assessment of which wolf was involved would enhance the relia-
bility. This means to enhance identification through means other than DNA, such as
registering visual or behavioral characteristics that describe the individual wolf, similar
to the registration of individual wolves in Yellowstone park [97]. Another possibility is to
use the howl as means to identify individual wolves [98], although loners tend to howl
less than members of a territorial pack [99].
Of the 17 identified loners, three loners (nr 2, 3, and 13) were not associated with
kills of farm animals (Table 2). Two of them were seen (wolves nr 2 and 3). Thus, the
probability of actually observing a loner that could not be detected by farm animal kills
was at least 2 out of 17, so 12%. These two wolves could be the tip of an iceberg and
representatives of a larger subpopulation of unobserved wolves: wolves that did enter
the Netherlands, but did not kill farm animals and went unobserved. If so, it is remarka-
ble that a subpopulation of loners enters and crosses the Netherlands without killing
farm animals; sheep in particular, as sheep are considered easy prey [8,34–36]. In this
respect, there are three significant observations. First, the three wolves mentioned above
were present in the same or similar areas where other wolves have killed sheep. Second,
the sheep density in the Netherlands is three times higher than in German states where
the parent packs live [100]. Therefore, the Dutch conditions make it even more unlikely
that a loner does not encounter and kill a sheep. Thirdly, access to sheep has hardly any
threshold, as by the absence of wolves over 170 years, Dutch sheep are kept without
special protection from wolves. Thus, the availability of sheep cannot explain why there
are loners that do not kill sheep. Other researchers also showed that prey preference
cannot be fully explained by availability [8,35,101–106]. Therefore, prey preference is
apparently not merely a matter of stochastic laws.
Other explanations for selecting sheep as prey are also mentioned, such as devel-
opment of specialization [101,107,108], or the development of an ecotype with a feeding
preference or a feeding habit [108,109]. Thus, learning mechanisms are involved. Refer-
ences to learning mechanisms are “development of hunting experience” and “learning
from parents” [10,13,35] or “learning differences between packs” [102]. Indeed, similarity
between choices of wolves in the parent pack and their migrating offspring gives a cor-
responding explanation. This possibly applies to the three wolves mentioned above, if
they stuck to prey familiarized to them through experiences gained in the parent pack.
The loners were young wolves at detection in the Netherlands with a median age of 20
months, as commonly found for young dispersing wolves [3,42]. If in the period before
dispersal, parent packs fed their young wild prey only and showed their maturing young
how to find, approach, and kill wild prey (like roe deer), then the loner has left the pack
only familiarized with these species. In cases where sheep have not been part of the diet
or hunt, they have not learnt about sheep as prey. We suggest that a wolf will have no
drive to explore new prey, such as sheep, as long as familiar wild prey density sustains
survival and reproduction. The common notion that sheep are easy prey may only be
true for wolves that are familiar with sheep. Conversely, the prediction that wolves will
not kill sheep if there is sufficient wildlife [35,36,110] may be incorrect for wolves that are
familiar with sheep.
Wolf nr 3 is a special case, as this wolf was not linked to the killing of sheep, but his
parent pack was. In the reasoning above, this is not expected. Wolf nr 3 was born in 2015
(Table 1). There were no sheep kills in or near his parent pack territory in 2015. However,
the first known kill of sheep was in 2016 as DNA samples showed that the parents
(GW203f and GW339m) killed sheep together on 24 January 2016 [52]. Wolf nr 3 would
then have been 8 months old. It is not clear if at that time he was still present in the pack,
as his DNA was never detected at sheep kills in or near his parent pack territory. Thus, it
is possible that wolf nr 3 had already left the pack early in January 2016 and had not
gained experience in sheep killing in the parent pack. Leaving the pack at a relatively
young age in January or February probably was also the case for wolves nr 1, 6, and 7,
which were 9 to 10 months old when they reached the Netherlands (Table 1). However,
Animals 2021, 11, 1801 13 of 20
the conclusion in the case of wolf nr 3 is that parent pack and loner behavior in killing
sheep is dissimilar.
In cases where the abundance of familiar wildlife prey is falling, it cannot be ruled
out that a naïve wolf starts exploring farm animals like sheep on its own. Roe deer are
wild ungulates strongly preferred by German wolves, making up 52–54% of the con-
sumed biomass [34,111]. Therefore, it could be expected that German wolves, familiar
with and normally focused on roe deer, tend to start to kill sheep in areas where sheep
densities are high and roe deer density falls. As discussed above, the Netherlands has a
sheep density three times higher than the German states where the parent packs are liv-
ing. However, the Dutch roe deer density of 3.7/km2 is likely to be lower than densities in
the German states where the parent packs are living. For instance, in Sachsen the esti-
mated density was 4.8 in 2015, as calculated from the annual roe deer cull [112], wolf kills
[34] and agriculture and nature area size [113]. The relatively high sheep density and
likely low roe deer density in the Netherlands make it all the more remarkable that loners
2, 3, and 13 did not start to kill sheep when entering this country. It strengthens the con-
clusion that prey preference of loners cannot be fully explained by prey availability and
that early life experiences in the parent pack are involved. Exploration of new sources is
seen with wolf nr 13, which did not kill sheep during migration (Table 2) but did kill
sheep once after settlement in February 2019 as a single wolf. This was in October 2019
[32]. It is not known however what triggered this kill. A trigger could be a drop in wild-
life abundance at that time or an increase of disturbances of wildlife by tourists or
shooters [12].
Three of the 17 identified loners (nr 1, 6, and 7) were in close proximity to humans
(Table 2). An explanation could be that the Netherlands is densely populated with 513
people per km2 [114], which may result in proximity. However, the wolves travelled
through human settlements during daylight and did not try to avoid the human envi-
ronment (see references in Table 2). Even in the Netherlands, and especially in villages
and towns where wolves have been seen travelling, there is ample opportunity to travel
through forests or fields adjacent to these human settlements rather than through streets
with traffic, pedestrians, and bicycles, like in the cases of wolves nr 1 and 6 (Figures 1 and
2). The simplest explanation would be then that these wolves have already had positive
reinforcement and experience with a human environment and chose this over adjacent
environment at distance from humans.
Positive experience with a human environment may have started in the parent pack.
Indeed, parent pack members that were known to be in close proximity to humans or
human settlements had similarly behaving loners (Table 4). Moreover, similarities in
proximity to humans applied to siblings. Both wolf nr 1 and his one-year-younger
brother GW369m, which stayed in Germany [50], and the two brothers nr 6 and 7 were in
proximity to humans. In addition, if there were no proximity references, then this also
applied for both, siblings and parent packs, as in the case of half-brother nr 4 and
half-sister nr 11 and sisters nr 2 and 17 (Tables 2–4). These findings confirm that besides
prey choice, young wolves may learn by following and imitating their parents or siblings
[4,9] to be in proximity of people. Already in the parent pack, young wolves may expe-
rience that human environments and humans need not be feared and may even be re-
warding. The consistency between parent pack and loner choices regarding proximity to
the human habitat corresponds to other studies where habitat preference of dispersing
wolves is shown to be determined by early life experiences [25,26]. Such vertical trans-
mission of preferences is thought functional for survival [4,17–20]. However, this trans-
mission implies that human–wolf conflicts may increase not only because of expanding
wolf populations, but also by reproduction of parent packs with the habit to be in
proximity of humans and a subsequent dispersal of a new generation that also prefers to
be in proximity to humans.
Kuijper et al. [36] recommend discouraging wolves from approaching human habi-
tat by deterring them while maintaining high abundance of wildlife prey. They follow
Animals 2021, 11, 1801 14 of 20
Huber et al. [115] and Newsome et al. [116], who assume that European wolves are ha-
bituated to humans, though it is not clear what “habituation” means in these references.
Moreover, Kuijper et al. [36] do not refer to rearing conditions of wolves and their con-
sequences for the development of a tendency to approach human habitat. LCIE [24] as-
sumes positive conditioning as part of wolf habituation, through food, dogs, or play
opportunities. However, LCIE only recommends deterring of so-called bold wolves.
These wolves would repeatedly tolerate or actively approach people within 30 m. The
main tool would be to prevent people from feeding wolves. Again, recommendations do
not clearly link with the parent pack, though LCIE does recognize maternal and sibling
influences on the development and consistency of bold wolf behavior. For unknown
reasons LCIE does not mention paternal influences.
Given the literature about vertical transmission of experiences as mentioned before
and the results in this study, we presume that prevention of problems may be promoted
by discouragement of parent pack wolves approaching human dwellings or humans. A
deterring focused at the reproducing pack is also expected to increase the threshold for
young inexperienced wolves to follow and copy the choice to enter the human habitat. As
a consequence, a tendency to find rest, shelter, and food in environments outside human
dwellings may be maintained across generations. Moreover, deterring bold wolves only
would be a very late measure. In our view the deterring should take place as soon as the
approach happens, so with any wolf that crosses villages or approaches humans, whether
fitting the definition of “bold” or not. This is in contrast to the currently accepted policy
in Germany, the Netherlands, and Belgium not to take measures when a non-bold wolf
enters a village. A spin-off of keeping wolves at a distance from human settlements could
also be that wolves are less likely to encounter and subsequently gain experience with
domestic animals.
When weighing the problem of proximity to humans with the problem of killing
farm animals, the latter is more severe, as a wolf attack has a severe impact on the welfare
of involved sheep, results in production losses [117] and it may lead to (illegal) killing of
wolves [35,36,100,118,119]. Moreover, it is a particular problem with loners as their mi-
gration pattern is difficult to predict. As a consequence, prevention measures may be
taken ad hoc, too late, and only after kills of farm animals have taken place. In addition,
parent packs may have learnt to overcome prevention measures, such as electric fences
[106], for instance by jumping over them. Young wolves may therefore have also learnt to
overcome (electric) fences. So when migrating, such wolves may not be sufficiently
thwarted to kill farm animals. The difficulty in predicting the occurrence of problems
with migrating wolves implies the need to reduce their likelihood in another way, which
can be through developing methods preventing farm animal killing experiences, while
the individual wolf is still in the parent pack. Thus, in particular, herds near the pack
should be properly protected. However, current methods such as electric fences and
guarding dogs are not always effective or possible [106]. Therefore, additional methods
need to be developed. We suggest testing the application of a sensor in two sheep in a
fenced herd, wirelessly connected to a deterring system that can cover and protect the
herd. This sensor would be immediately activated by the stress in a sheep due to the ap-
proach of a wolf [120]. A second measure that can teach wolves not to select sheep is a
collar for sheep, activated the moment a wolf bites in the neck and then delivering a
painful electric shock only to the wolf [121]. A third way is to provide a sheep carcass
injected with an emetic, placed near the wolves’ den. When an adult wolf feeds on this
cadaver, the nausea is expected to stop the wolf from selecting this carcass and possibly
other sheep as food, also in the course of feeding the young. The methods combined
contribute to teaching wolves and their offspring not to approach, bite, and consume
sheep. This seems effective when alternative food resources are abundant, such as wild
ungulates. The problem of (unpredictable) attacks on sheep may therefore be largely
solved, if all levels of the wolf’s hunting (the approach, the biting, and the consumption)
are negatively reinforced in parent packs. Those packs would thus raise new generations
Animals 2021, 11, 1801 15 of 20
of migrating wolves without prior experience with sheep as prey and not necessarily
killing sheep.
5. Conclusions
Though with a limited sample size, this study shows that the behavior of parent
packs significantly predicts the behavior of their migrating offspring. Parent packs that
killed sheep and were or were not regularly seen near humans or human settlements had
similarly behaving loners. In addition, there were loners that behaved unproblematically
like their parent packs: they did not kill sheep nor were they near humans. This suggests
that the common notion that sheep are easy prey may only be true for wolves that are
familiar with sheep. Conversely, the prediction that wolves will not kill sheep if there is
sufficient wildlife may be incorrect for wolves that are familiar with sheep. The results
confirm that in addition to prey choice, young wolves may learn to be around humans by
following and imitating their parents or siblings. Vertical transmission of experience im-
plies that human–wolf conflicts may increase not only due to expanding wolf popula-
tions. They may also increase by reproduction of parent packs with a habit of killing
sheep or being near humans, and a subsequent dispersal of a new generation with similar
habits. The difficulty in predicting the occurrence of problems with migrating wolves
implies the need to reduce problems by new methods. These new methods should gen-
erate negative experiences of killing farm animals and being in proximity to humans
while the individual wolf is still in the parent pack.
Author Contributions: Conceptualization, D.v.L., N.S., and D.J.; methodology, D.v.L., N.S., and
D.J.; validation, D.v.L., N.S., D.J., and P.M.; investigation, D.v.L., N.S., and D.J.; writing—original
draft preparation, D.v.L., N.S., and D.J.; writing—review and editing, D.v.L., N.S., D.J., and P.M.;
supervision, D.J. and P.M. All authors have read and agreed to the submitted version of the man-
Funding: The publishing was partially funded by the AnimalWise foundation.
Institutional Review Board Statement: Not applicable.
Data Availability Statement: The data are contained within the article.
Acknowledgments: We would like to thank Sebastian Collet of the Conservation Genetics Group
of the German Senckenberg Forschungsinstitut, Konstantin Schanze of the Saxon State Office for
Environment, Agriculture and Geology, Germany, Verena Harms of the Landesamt für Umwelt,
department Naturschutz und Brandenburger Naturlandschaften in Brandenburg, Germany and
Arjen de Groot at the Ecological Genetics and Ecosystem Services of Wageningen Environmental
Research, University of Wageningen, Netherlands, for their cooperation in providing wolf data.
Conflicts of Interest: The authors declare no conflict of interest. N.S., D.J., and D.v.L. are co-author
of patent SI 25485 A that describes a collar for sheep to protect them against wolf attacks. These
facts had no influence on the content of this manuscript.
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Within the 2700km² Beltrami Island State Forest, near the W edge of the primary range of Canis lupus in Minnesota, wolf population density was low at the start of the study in 1972 but increased substantially up to 1977 (end of study). At least 8 of 13 social units present in mid-1976 had formed since 1972. Size of litters of established packs averaged 4.6 pups, and those of newly-formed pairs averaged 4.1. Mortality decreased over the study period, and recruitment of young wolves exceeded mortality following legal protection. A high rate of dispersal of young from packs was documented. Dispersal peaked in autumn. Most wolves paired within a few days of leaving their packs. Average territory size decreased as both population and pack numbers increased. Behaviour of alpha males, alpha females and subordinate members of the packs is discussed. Deer and moose comprised 94% of animal biomass eaten by wolves, with deer along accounting for 67%. Seasonal differences in food taken and energy requirements are noted.-P.J.Jarvis
This pilot study aimed to deliver initial findings about the territorial and dispersal behaviour of wolves in Germany. Six wolves (2 adults, 4 pups) were fitted with GPS-GSM collars. The average territory size of adult wolves was 203 km² MCP 95 and 328 km² MCP 100. During the study period 2 out of 4 subadult wolves wearing a radio collar dispersed from their natal territory. A young male left its natal pack when 12 month old and dispersed 1 500 km (800 km straight line) all the way from Lusatia to Belarus. His radio-collared brother made smaller excursions around his natal territory before establishing an own territory bordering his parents' territory when 22 months old. During their excursions both wolves crossed fenced highways. In the Lusatia Wolf area the percentage of forest cover is 40 %, which is higher than the national average (30 %). Within their territories wolves used forest as available, preferred areas with heather and succession but avoided fields and pastures. The results of this pilot study highlight the enormous adaptability of wolves, underscoring how small our country is from a Wolf's perspective.