Access to this full-text is provided by Pensoft Publishers.
Content available from Nature Conservation
This content is subject to copyright. Terms and conditions apply.
Dogs, not wolves, most likely to have caused the death
of a British tourist in northern Greece
Yorgos Iliopoulos1, Christos Astaras2, Eirini Chatzimichail1
1 Callisto Wildlife and Nature Conservation Society, Mitropoleos 123, GR-54621, essaloniki, Greece
2Forest Research Institute, ELGO-DIMITRA, GR-57006 Vassilika, essaloniki, Greece
Corresponding author: Yorgos Iliopoulos (yiliop2@gmail.com)
Academic editor: A. Grimm-Seyfarth|Received 9 February 2022|Accepted 24 September 2022|Published 20 October 2022
https://zoobank.org/17DA017D-DFA9-46C3-AC60-A581E4D080AF
Citation: Iliopoulos Y, Astaras C, Chatzimichail E (2022) Dogs, not wolves, most likely to have caused the death of a British
tourist in northern Greece. Nature Conservation 50: 115–143. https://doi.org/10.3897/natureconservation.50.81915
Abstract
Wolf (Canis lupus) populations have recovered and expanded across many parts of the world thanks to
conservation eorts, including improved legal status and restoration of their prey. Concurrently, public
concerns regarding the risk of wolf attacks on humans and livestock are increasing as wolves occupy human-
dominated landscapes. We examined a unique case in Europe allegedly involving wolves in the death of a
female British tourist, aged 64, in northern Greece in September 2017. is incident received extensive
international media attention and yet many fundamental details of the case area are lacking, including
whether local livestock guarding dogs played a role. To assist in resolving the case, we conducted an extensive
literature review which documented 13 criteria linked to the risk of either a wolf and/or a dog attacking a
human. We also conducted a camera trap survey (October to December 2017) soon after the fatal attack to
calculate the activity overlap among humans, dogs and wolves. Sucient data were available for assessing
11 of the 13 criteria. For the remaining two, the required data were either not analysed (i.e. canid DNA
collected from the attack site), not appropriately collected (i.e. DNA from the mouths of suspected dogs)
or were collected, but misinterpreted (i.e. the post-consumption patterns of the victim’s corpse). Via this
combination of evidence, we conclude that this case involved a fatal dog attack. is assertion is supported
by evidence such as the: a) high dog-human activity overlap at the attack site which peaked during the attack
time as opposed to near zero wolf-human activity overlap at the same time, b) presence of a large pack of un-
supervised dogs, c) high ratio of male dogs in the dog pack, d) close vicinity of the attack site to dog owner’s
property and e) previous documented aggression of these dogs towards humans. e consumption patterns,
time scale and location of the victim’s remains indicate a posthumous consumption of the corpse possibly by
the same dogs and/or by wild scavengers including wolves. A multidisciplinary approach, such as this one, in
the assessment of putative wildlife attacks on humans can reduce misidentications of the responsible species
by forensic authorities and, therefore, prevent unfounded decrease in public tolerance for large carnivores.
Nature Conservation 50: 115–143 (2022)
doi: 10.3897/natureconservation.50.81915
https://natureconservation.pensoft.net
Copyright Yorgos Iliopoulos et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
RESEARCH ARTICLE
Launched to accelerate biodiversity conservation
A peer-reviewed open-access journal
Yorgos Iliopoulos et al. / Nature Conservation 50: 115–143 (2022)
116
Keywords
animal attacks on humans, forensic analysis, large carnivores, livestock guarding dogs, multidisciplinary
approach, wolf
Introduction
e wolf (Canis lupus L.) has expanded across many parts of the world over the last
three decades, thanks to conservation eorts related to its legal protection and restora-
tion of its habitats (Chapron et al. 2014; Cimatti et al. 2021). is recovery of wolf
populations, although often heralded as important for restoring ecological processes
(Ripple and Beschta 2012; Boyce 2018), is not without risk, including the potential of
increasing human-wildlife conict due to wolf depredation of livestock and game ani-
mals (Janeiro-Otero et al. 2020). Moreover, concerns about the risk that wolves pose to
human safety are gaining public attention (Linnell et al. 2003), as wolves increasingly
occupy human-dominated landscapes (Kuijper et al. 2019).
Despite the fact that there are more than 17,000 wolves in Europe (Boitani 2018)
and 75,000 in north America, with presence also in landscapes which are home to mil-
lions of people, reports of wolves attacking humans since the early 20th century are rare
(Penteriani et al. 2016; Linnell et al. 2021). Wolf attacks resulting in human injuries or
fatalities have been reported in published literature, unpublished/historical reports and
the media (Linnell et al. 2002; Linnell et al. 2021). In general, wolf attacks on humans
can be categorised as: 1) attacks by unhealthy/injured wolves, 2) provoked or defensive
attacks and 3) predatory attacks (Linnell et al. 2021). Attacks by unhealthy wolves have
been reported mainly in areas with rabies prevalence in wildlife, like India (Isloor et al.
2014), China (Wang et al. 2014), Iran (Gholami et al. 2014), Turkey (Turkmen et al.
2012; Ambarli 2019), Russia (Sidorov et al. 2010), as well as other countries from the
Middle East, Eurasia and Asia (Linnell et al. 2021). Provoked wolf attacks on humans
are rarer and have involved cases where wolves injured humans in defence of their life,
prey or ospring (Linnell et al. 2021). Predatory attacks were mainly reported from
areas with low natural wolf prey availability, wolf habituation with anthropogenic food
sources, such as livestock, oal remains and garbage, high human density in rural set-
tings and absence of rearms (McNay 2002; Löe and Röskaft 2004; Lescureux and
Linnell 2014). Such conditions are mostly found in the Middle East and Asia, includ-
ing India (Jhala and Sharma 1997; Rajpurohit 1999), Iran (Behdarvand et al. 2014;
Behdarvand and Kaboli 2015) and Israel (Linnell et al. 2021). Predatory attacks on
humans by healthy wolves have been also reported in Europe and North America in the
20th and 21st century (Linnell et al. 2002; Mc Nay 2002; McNay and Mooney 2005;
McNay 2007; Butler et al. 2011; Penteriani et al. 2017; Linnell et al. 2021; Nowak et
al. 2021). However, in the last 40 years, since scientic studies on wolves have been
carried out, only two people have been killed by wolves in North America, while in
western Europe (excluding Russia and some neighbouring countries where rabies is still
prevalent), no wolf predatory attack on humans has been veried (Linnell et al. 2021).
No evidence of wolf involvement in a human fatality in Greece 117
As reports of wolf attacks on humans and increased livestock depredation may im-
pede species conservation eorts (Linnell et al. 2021), great care is needed in assessing at-
tacks putatively attributed to the species. Reports of wild predator attacks on humans and
livestock – especially when receiving wide media coverage – have the potential to dispro-
portionately decrease public tolerance towards those species, with broader repercussions
for wildlife conservation (Kansky and Knight 2014; Homann et al. 2017; Penteriani et
al. 2017; Bombieri et al. 2018). Unsubstantiated incidents of human-wolf conict, in
particular involving human injuries or fatalities, can increase public fear of wolves. Fear
has been used by individuals or interest groups to promote public dislike for wolves and
to reduce the legislative protection aorded to the species (Linnell and Alleau 2015).
We examine here the 2017 case study of an alleged wolf involvement in the fatal
attack of a female British tourist, aged 64, in northern Greece, because of the potential
ramications it could have for wolf public perception across Europe, especially given
the extensive media attention it received both nationally and internationally (Arbieu et
al. 2021). Specically, we tested the hypothesis that wolves were indeed responsible for
the fatality by examining factors associated with: a) wolf and dog attacks on humans
reported in literature, b) relying on case-related evidence from ocial reports and c)
our own eld investigations, initiated soon after the attack, regarding wolf and dog
occurrence and activity patterns at the attack site and adjacent areas. e criteria and
assessment protocols proposed and used may help reduce the chance of predator misi-
dentication in future human attack cases.
Case report
According to the ocial police report to the District Attorney, on 21 September 2017,
the victim called her family in Britain at about 17:00 h Greek local time (GMT+2),
which was still daylight, to report that she was being “attacked by erce dogs” along the
Petrota-Maroneia section of the E6 European long distance tourist trail in Rhodope
Prefecture, northern Greece. Local witnesses, including two Austrian tourists whose
written report to WWF Greece was shared to us with their consent, conrmed hav-
ing seen her one hour earlier (4 pm) leaving Petrota Beach (~ 3.7 km from the attack
site) heading on foot towards Maroneia Village along the car-accessible dirt/forest road
which is part of the E6 European long-distance hiking trail (Figs 1–2). In response,
the re brigade initiated, that same evening, a missing person search in the area, but
returned without success. e next morning (22 September 2017), the search resumed
with additional personnel from the local police department, a special search and rescue
unit of the re brigade and local volunteers, continuing through the night and beyond.
Eventually, using tracking dogs, some personal belongings (incl. passport, clothing)
and bodily remains of the victim were discovered on the morning of 23 September
2017, approximately 36 hours after the initial distress call. e collection of additional
remains concluded 72 h following the attack. e remains were dispersed along an ap-
proximately 15,000 m2 area adjacent to the E6 trail/road, 3.7 km from Petrota Beach
– or approximately 1 hour of leisure walk (Figs 1–2). According to the ocial police
Yorgos Iliopoulos et al. / Nature Conservation 50: 115–143 (2022)
118
Figure 1. Area delimited by solid line indicates the approximate boundaries of the study area, the dashed
line indicates the approximate location of the local wolf pack rendezvous site. Star symbols indicate cam-
era traps deployed at the “Attack area”, while closed square symbols indicate location of camera traps
deployed in the “Broader area”.
Figure 2. Location of the attack site in relation to a nearby goat herd corral and watering spot.
No evidence of wolf involvement in a human fatality in Greece 119
report, the attack site (60 m a.s.l.) was determined to be approximately 40 m from the
forest road, 200 m from an actively used seasonal night-time corral for a 750-strong
goat herd and 90 m from the herd’s watering troughs (Figs 2–4). e herd was pro-
tected by at least ten free-roaming livestock guarding dogs (Figs 5–6).
According to the Coroner’s report, the tourist’s body was almost completely con-
sumed with only ten bone fragments (parts of skull, lower jaw, femur and tibia) and
soft tissue (both lungs, part of the heart and small parts of skin) retrieved by the police.
All remains had signs of animal bites and/or consumption. Amongst the retrieved be-
longings were torn clothing (incl. jacket, trousers, shirt, hat) with traces of dry blood.
Initial national and international media reported stray dogs or livestock guard-
ing dogs as being responsible for the death of the victim (e.g. “e Guardian” 2017,
“Express” 2017). However, a 26 September 2017 article in e Times (London) (de
Bruxelles and Carassava 2017) cited the Coroner’s belief, prior to the completion of the
laboratory analysis of remains, that the victim “may have been attacked by other wild
animals, like rabid wolves” judging by the state of the victim’s remains. is statement,
which was widely reproduced by national and international media, contributed to an
already ongoing debate on the future of the protection status of wolves in European
countries, including a call by the German Federal Ministry of Agriculture, just four days
after e Times’ article, to regulate their national wolf population by hunting (Heine
2017). Recognising the potential long-term impact of this case on public perceptions of
human-wolf interactions in Europe, we initiated this study on 2 October 2017.
Figure 3.e attack site over the E6 trail (forest road passable for vehicles).
Yorgos Iliopoulos et al. / Nature Conservation 50: 115–143 (2022)
120
Study area
Our study area extended over 21 km2 in the Prefecture of Rodope in northern Greece
(Fig. 1). It was dened as the area encompassing: (a) the attack site, (b) the grazing area
of the goat herd kept at the corral adjacent to the attack site and (c) potential nearby
wolf pack home sites (i.e. rendezvous sites) as dened by habitat modelling (Iliopoulos
et al. 2014) because wolf presence in the area was unknown at the onset of the study.
Figure 4.Watering spot of the goat herd corral adjacent to the attack site at 90 m, occupied throughout
the study duration and until early November 2017 where remains of the victim were also retrieved.
No evidence of wolf involvement in a human fatality in Greece 121
Figures 5, 6. Unsupervised livestock guarding dog images from the goat ock occupying the corral
adjacent to the attack site.
6
5
e study area ranges from 0–612 m a.s.l and is characterised by dense evergreen
Mediterranean shrubland dominated by Quercus coccifera. Cultivated and abandoned
olive groves extend across parts of the lower elevation and coastal areas. In addition to
the goat herd that are grazed in this area, there is one more free-ranging livestock herd
Yorgos Iliopoulos et al. / Nature Conservation 50: 115–143 (2022)
122
within the study area and ve around the periphery (Fig. 1). e dirt forest road that
forms the E6 European long-distance hiking trail traverses the study area parallel to
the coast (Figs 2–3).
Methods
Our study was complementary to the on-going police investigation at the time and
tasked under a permit issued by the Hellenic Ministry of Environment and Energy
with: (a) assessing wolf and dog presence and activity patterns in the study area and
(b) providing an expert wildlife opinion on the case. All material (e.g. torn clothes,
human bones, animal hair and faeces) collected at the attack site, which could be used
to extract genetic material for DNA analysis of the involved predator, were held by
the police department and were not accessible for external analysis. Nevertheless, in
March 2019, the District Attorney handling the case granted access to the police and
coronary-related reports and, hence, we included their ndings in our analysis.
Reviewing both canid ecology literature and forensic cases involving large carnivore
and dog attacks on humans, we identied a list of criteria and factors either linked to
the risk of a human being attacked or useful for distinguishing the responsible predator
(Table 1). We then used these criteria to assess our case. is approach was deemed nec-
essary because the full consumption of the victim’s corpse and the time it took to locate
the victim’s remains severely compromised the collection of evidence related to the attack
for predator identication, such as size and shape of bite marks, location of injuries to
the soft tissue corpse (i.e. Fonseca and Palacios 2012; Fonseca et al. 2015) or analysis
of saliva DNA from fresh body wounds collected soon after the attack and prior to any
post-mortem consumption (i.e. Caniglia et al. 2013; Caniglia et al. 2016). Furthermore,
multi-criteria assessment was also needed due to an additional complication in our case;
the inability to determine if the biological samples (hair, faeces) collected at the attack site
were from the predator or subsequent scavengers, because it took 36–72 h from the initial
distress phone call to locate all the victim’s remains. Even though these samples were not
ultimately analysed for canid DNA, the interpretation of such analysis would have been
problematic since scavenging could have taken place prior to collection. Other authors
have also urged to not depend solely on forensic pathologist reports in cases of animal-
related human fatalities, but to include ecological and environmental characteristics of
the attack as well (i.e. Fegan-Earl 2005; Shields et al. 2009; Fonseca and Palacios 2012).
Table 1. Criteria and factors used in the present study to assist in identifying the species (wolf or dog)
most likely to be responsible for the attack.
Criteria and Factors for identication of predator
Description and Rationale (species applicable) [Source] Data collection method
1. Encounter rate between carnivore and human (Dog, Wolf)
Probability of an attack increases with increased encounter rate. camera trapping
[Penteriani et al. 2016; Penteriani et al. 2017; Martin et al. 2018]
No evidence of wolf involvement in a human fatality in Greece 123
Criteria and Factors for identication of predator
Description and Rationale (species applicable) [Source] Data collection method
2. Location of attack site in relation to carnivore territory (Dog, Wolf)
Carnivore aggression towards humans may be escalated closer to certain areas of their territory. Most dog
attacks, causing injury or death on humans are connected to their owner’s property. Wolves may show
aggression to humans close to home-sites and denning areas.
police reports, camera trapping,
eld observations, wolf home-
site predictive model
[Rubin and Beck 1982; Borchelt et al. 1983; McNay 2002; Patronek et al. 2013; Notari et al. 2020].
3. Carnivore health / body condition (Dog, Wolf)
Rabid wolves and dogs attack all mammals, including humans when rabies is endemic with no consumption
of the victims [Turkmen et al. 2012; Ambarli 2019]. Rabies in Greece was not present in the country during
the study period. Wild carnivores with compromised health (i.e. old age, injury, disease) may be more prone
to seek food close to humans or identify humans as prey and attack.
national rabies reports, camera
trapping and eld observations
[Penteriani et al. 2016; Nowak et al. 2021]. Healthy wolves may also attack and kill humans. [Butler et al
2011; Behdarvand et al 2014; Behdarvand and Kaboli 2015]
4. Group size (Dog)
Dog group size is positively related to the probability of a fatal attack on humans. A single aggressive move
towards a human by one dog can trigger the attack of the rest of the pack, escalating the severity of the attack
often until the victim is immobilised or dead.
camera trapping, eld
observations
[Borchelt et al. 1983; Kneafsey and Condon 1995; Raghavan 2008; Santoro et al. 2011; Patronek et al. 2013]
5. Dog(s) body size (Dog)
e presence of large-bodied dogs may be related to more frequent and severe attacks on humans. camera trapping, eld
observations
[Roll and Unshelm 1997; Mikkola et al. 2021]
6. Number of male dogs (Dog)
e number of male unneutered dogs present in a group is positively related to the probability of an attack on
humans. Most fatal dog attacks on humans involved male dogs.
camera trapping, eld
observations
[Shuler et al. 2008; Hsu and Sun 2010; Patronek et al. 2013; Matos et al. 2015; Notari et al. 2020; Mikkola
et al. 2021]
7. Wolf prey availability and wolf habituation (Wolf)
Low natural prey availability and/or human-related food provision may aect the frequency with which
wolves visit human settlements in search of food, leading to fearless behaviour towards humans and may
predispose them on predatory attacks in any location of their territories
camera trapping, eld
observations, mapping of
human related food resources
(i.e. livestock herds)
[Sidorovich et al 2003; Heilhecker et al. 2007; Behdarvand and Kaboli 2015; Nowak et al. 2021]
8. Season of attack related to wolf biological cycle (Wolf )
Wolf attacks on humans are mostly recorded between May and August Police reports
[Rajpurohit 1999; Behdarvand and Kaboli 2015]
9. Dog socialisation to humans and previous aggression (Dog)
Lack of, or negative, socialisation and/or previous aggression of dogs to humans may increase probability of
a dog attack.
camera trapping, eld
observations, witness
testimonials
[Patronek et al. 2013; Marion et al. 2018; Mikkola et al. 2021]
10. Human supervision (Dog)
Lack of human supervision of free roaming dogs increases the chance of an escalated in severity attack and,
therefore, the probability of its being fatal.
camera trapping
[Rubin and Beck 1982; Borchelt et al. 1983; Patronek et al. 2013]
11. Bite patterns (Dog, Wolf)
Location and distribution of bites over the victim’s body may reveal species and individuals involved. police and coroner reports
[Santoro et al. 2011; Fonseca et al. 2015]
12. Post-mortem consumption patterns and rates (Dog, Wolf)
Extent, rate and patterns of post-mortem consumption may be indicative of the carnivore species involved
and its group size. Attacks and post-mortem consumption of humans by dogs is not typically driven by
hunger, although cases of human corpses being consumed almost completely, even of their owners, have been
reported. Dog consumption patterns on human resemble those of wild canids.
police and coroner reports
[Borchelt et al. 1983; Haglund et al. 1989; Rothschild and Schneider 1997; Avis 1999; Wilmers and Stahler
2002; Peterson and Ciucci 2003; Christiansen and Wroe 2007; Steadman and Worne 2007; Buschmann et al.
2011; Fonseca and Palacios 2013; Behdarvand and Kaboli 2015; Fonseca et al. 2015]
13. Sampling of genetic material from bites and wounds
DNA obtained from samples collected from the mortal remains can be used to identify responsible
carnivore(s) for the attack and/or post-mortem consumption.
coroner and genetic laboratory
reports
[Sundqvist et al. 2007; Caniglia et al. 2013; Harms et al. 2015; Caniglia et al. 2016; Plumer et al. 2018;
López-Bao et al. 2017]
Yorgos Iliopoulos et al. / Nature Conservation 50: 115–143 (2022)
124
Data collection
Our eld data collection commenced on 2 October 2017, ten days after the body
remains of the victim were located and one week after the coroner’s interview in “e
Times”. It concluded two months later (5 December 2017) and after the goat herd and
its dogs had left the corral at the attack site (12 November 2017). While a wolf pack’s
movement pattern can change when the pups start to follow adults away from the den,
our eventual data showed this to have taken place in November. Our data collection,
therefore, commenced suciently close to the incident to contain relevant data about
dog and wolf activity in the area at the period of the attack.
While wolves are widely distributed in northern Greece, at the onset of the study,
the spatio-temporal presence of wolves proximate to the attack site was not known.
We deployed six camera traps (Reconyx RC60, Bushnell HD Trophy Cam) along for-
est roads and paths to: a) examine the presence of wolves during the study period and
their breeding status and population size and b) record whether and to what extent
wildlife, livestock, dog and human activities overlapped in time and space. Camera
traps have been widely used across the world to examine interspecies interactions,
including with humans (e.g. Muhly et al. 2011). With that in mind, we grouped the
cameras in two clusters of three cameras each, based on their distance from the site of
the attack. One cluster (“Attack area”) monitored animal and human activity within
the grazing area of the goat herd stationed at the corral adjacent to the attack site. Spe-
cically, one camera was placed on the E6 trail (dirt road) 20 m from the attack site,
a second one was placed 600 m to the west and the third 1,600 m to the northwest
along a trail/road leading to the E6 trail (Fig. 1). e cameras of the second cluster
(”Broader area”) were placed at locations closer to and in-between two areas identied
as potential wolf pack rendezvous sites (home-sites), based on criteria developed by Il-
iopoulos et al. (2014) that links water presence, distance from roads, forest cover and
human infrastructure (villages) with rendezvous site suitability in Greece. ey were
2,100 m, 3,800 m and 4,500 m (Euclidean distance) from the attack site (Fig. 1).
e camera traps use passive infrared sensors to detect heat and movement within
a funnel-shaped area in front of the camera (radius ~ 10 metres) and are silent. ey
were set to record three consecutive images per triggering event (rapid-re mode)
which is known to capture even fast-moving objects (e.g. animals, vehicles). e
cameras recorded around the clock, including at night with the use of infrared light
which was invisible to mammals (covert type 940 nm “no glow”). e recorded im-
ages were time and date stamped. To avoid either theft or vandalism, the cameras
were carefully concealed in bushes by an experienced camera-trap user (YI), while
maintaining direct line of sight of the road. As this was a study conducted with the
specic intent of obtaining information to assist the police investigation, the place-
ment of the camera traps was unknown to the local population. erefore, the activ-
ity data obtained for both humans and animals were unaected by the presence of
the cameras.
No evidence of wolf involvement in a human fatality in Greece 125
Data analysis
For each photographed wolf or dog, we recorded its sex, age (pup, juvenile, adult) and
body condition, when possible. In addition, dogs were classied in three categories
(livestock guarding, hunting and stray dogs), based on their morphological character-
istics and their association or not with livestock herds and human activities (shepherds,
hikers, hunters). Especially for livestock guarding dogs (LGDs), we individually identi-
ed them, based on their size, coat type and pattern, tail shape and the livestock herd
with which they were associated. Furthermore, we classied LGD detections in two
categories: supervised (i.e. accompanying herd and/or shepherd) and unsupervised (i.e.
moving alone). We categorised human detections into those within vehicles and those
on foot or bicycles. e latter group was further divided into researchers (our team –
excluded from analysis), recreationists (hikers, hunters, bicyclists) and shepherds.
We considered photographs of a given species or category to be a dierent detec-
tion event when separated by at least one hour. For each camera trap cluster, we cal-
culated the relative abundance index (RAI) of each category of humans and animals
mentioned above using the following formula: RAI = n * 100/TD, where n = number
of detection events and TD = number of trapping days of the cluster’s camera traps.
We estimated the coecient of activity overlap (Δ), as dened by Ridout and
Linkie (2009), between: a) unsupervised LGDs and human - recreationists, b) wolves
and human - recreationists and c) unsupervised LGDs and wolves, using the pack-
age “overlap” (Meredith and Ridout 2021) in the R statistical package (R Core Team
2019). Δ can take values between 0 (no activity overlap across 24 h) and 1 (perfect
overlap). We calculated the condence intervals of Δ using bootstrapping (n = 1,000).
For this analysis, when multiple individuals were captured within a single detection
event, each animal was counted individually as per Muhly et al. (2011).
Results
In total, our camera trapping eort was 293 trap days (mean 49 ± 15 SD, n = 6); 120 for
the “Attack area” and 173 for the “Broader area” (Table 2). Wolf presence was conrmed
via records at all six cameras, consisting of individuals of one reproductive pack (breeding
pair and four pups). In addition, we conrmed the presence of golden jackals (Canis
aureus), which is a potential scavenger for the case study (Table2). e maximum jackal
family group observed was two animals, with most detections recorded in the “Attack
area”. All jackal detections occurred after dusk and before dawn. e encounter rate of
natural wolf prey species was very low, with roe deer (Capreolus capreolus) not detected
at all and wild boar (Sus scrofa) detected only once in the “Broader area” (Table 2).
Specically, considering both the camera trap data and the eld observations, we were
able to obtain the following information regarding the presence, activity patterns and
overlap of wolves, livestock guarding dogs and humans at the two sites.
Yorgos Iliopoulos et al. / Nature Conservation 50: 115–143 (2022)
126
Vicinity of attack site – “Attack area”
We recorded 11 wolf detection events. All occurred during night-time with the activity
pattern of wolves at the “Attack area” peaking at midnight and prior to sunset (Fig. 7,
chart a). Until 11 November, wolf detections (n = 4) involved solely the two adults of
the pack. By the time the pups, born in spring 2017, appeared in the cameras, the cor-
ral adjacent to the attack site was not systematically used by the goat herd.
We recorded 56 detection events of humans, of which 34 were of shepherds. e rec-
reationists (hunters n = 5, hikers/cyclists n = 17) ranged in group size from 1–3 individu-
als. Most detections were around noon and peaked during afternoon (Fig. 7, chart c).
We recorded 69 LGD detection events and identied in total 11 LGD individuals
belonging to the goat herd stationed at the corral adjacent to the attack spot, of which
at least eight were males. All but two of the dog pack were of large-bodied breed in
good physical condition (Figs 5–6). Activity at the corral (vehicles, shepherds, herd,
LGDs) was recorded daily until approximately 12 November, when the herd moved
out of the coral. After this date, the presence of the LGDs and its herd was periodic. As
of early November, a cattle herd started appearing in the cameras accompanied by two
additional LGDs, accounting for six LGD detections only. Hunting dog detections
were rare in this site (n = 3). No feral dogs were detected at the “Attack area”.
From those 69 LGD detection events, 45 were of “unsupervised dogs” (i.e. with-
out livestock or humans/vehicles recorded at the given camera within 1 h). Supervised
LGDs accompanying the goat herd were mostly detected during morning hours (9 am
– 12 noon), (Fig. 7, chart e). In contrast, unsupervised LGD activity was multimodal,
with most detections occurring either between 9 am -10 am or 4 pm - 7 pm (Fig. 7,
chart g). e evening activity of the goat herd LGDs was also corroborated by the eld
team during the camera trap deployment at the “Attack area”. Specically, at 7 pm
(night-time) on 3 October 2017, while the livestock were corralled, the authors (YI,
EC) observed and heard the LGDs barking spread out over a radius of several hundred
meters from the corral.
Table 2. Activity measures for wildlife, livestock guarding dogs and humans at the “Attack area” and
“Broader area”, based on camera trapping, expressed as relative abundance indexes (RAI’s) and sum of all
individual detections.
Species / Category Relative Abundance Index (RAI) Sum of all individual detections
“Attack area” “Broader area” “Attack area” “Broader area”
Wolf 914 47 43
Golden jackal 32 547 10
Wild boar 00.6 01
Livestock guarding dogs (LGDs) - Total 58 9169 62
LGDs with herd/shepherd 20 583 33
LGDs unsupervised 38 686 29
Hunting dogs 2.5 18 338
Livestock (goats, cattle) 76.6 9.25 - -
Shepherds 29 448 7
Recreationists (hikers, hunters, bicyclists) 18 12 49 32
Vehicles 189 120 227 207
No evidence of wolf involvement in a human fatality in Greece 127
Livestock presence was pervasive throughout the study period. We recorded 92 detec-
tion events of the goat herd from the “Attack area” and one cattle herd. In addition, we
observed several livestock carcasses in various stages of consumption along the E6 trail.
eir cause of death could not be determined in all cases (e.g. disease, disposal, predation).
“Broader area”
e number of detection events of wolves in the “Broader area” was higher than at the
“Attack area”, (RAI 14 and 9, respectively; Table 2), which is not surprising considering
that the camera sites were closer to highly suitable habitat for wolf rendezvous sites.
Figure 7. Density plots showing the activity pattern of wolves at (a) “Attack area” (n = 47) and (b)
“Broader area” (n = 43), humans (excluding researchers and shepherds) at (c) “Attack area” (n = 49) and
(d) “Broader area” (n = 32), supervised large guarding dogs with the herd at (e) “Attack area” (n = 83) and
(f) “Broader area” (n = 33) and unsupervised large guarding dogs at (g) “Attack area” (n = 86) and (h)
“Broader area” (n = 29). e time of the original observations at the camera traps are displayed as black
ticks below the x axes. Sample sizes refer to the sum of all detections (Table 2).
Yorgos Iliopoulos et al. / Nature Conservation 50: 115–143 (2022)
128
e actual presence of a rendezvous site was veried by spontaneous pack howling,
including pups, heard on October 7 by the eld team. e wolf activity pattern in the
“Broader area” was cathemeral (i.e. without a particular pattern; Fig. 7, chart b) and
resembled the one of unsupervised LGDs at the “Attack area” (Fig. 7, chart g).
Overall human detection events were fewer in the “Broader area”, with shep-
herds being seven times less frequently detected than at the “Attack area” (RAI 4
and 29, respectively). Of the recreationists, most events involved hunters (n = 16),
with hikers/cyclists detected only three times. is explains the human activity pat-
tern peaks at 7am and 2 pm – times when hunters go and return from their trips
(Fig.7, chart d).
e detections of LGDs in the “Broader area” was six-fold lower than at the “At-
tack area” (RAI 9 and 58, respectively; Table 2), and involved dogs from four neighbor-
ing farms (range of distance to cameras 1.4–2.6 km) and not the goat herd corralled
next to the attack site. e supervised and unsupervised LGD detections were almost
evenly split (9 and 10 detections, respectively). For both LGD categories, the activity
pattern was concentrated during daylight hours, peaked around noon and did not dif-
fer signicantly as in the case of the “Attack area” (Fig. 7, charts g and h). Hunting dogs
were seven times more frequently encountered than at the “Attack area” (Table 2). In
addition, we detected three suspected feral dogs. Livestock were also seven times less
frequently detected than at the “Attack area” (RAI 9.25 and 76.6, respectively; Table2)
and involved primarily cattle and goat herds other than the one at the “Attack area”.
Activity overlap
e activity overlap between wolves and humans (hikers, cyclists, hunters) at the “At-
tack area”, where the fatal attack took place, was signicantly lower than the overlap
between unsupervised LGDs and humans (0.04 vs. 0.55; Table 3). At 5 pm, which
is the time of the attack, the probability of wolves encountering humans during our
study period was near zero (Fig. 8, chart a). On the contrary, the overlap of unsuper-
vised LGDs and humans was at its peak at that same time (Fig. 8, chart b).
e very low activity overlap between wolves and humans was observed only at the
“Attack area” and not in the “Broader area”; the latter of which represents the activity
at the broader landscape beyond the immediate vicinity of the attack site (Table 3).
Moreover, activity overlap of LGDs by wolves at the “Attack area” was half of that in
the “Broader area” during the study period.
Table 3. Activity overlaps (Δ) between humans, wolves and LGD’s at the “Attack area” and “Broader area”.
Activity overlap pairs Overlap coecient Δ (95% CI)
“Attack area” “Broader area”
LGDs (unsupervised) & Humans 0.55 (0.42–0.68) 0.72 (0.56–0.89)
Wolves & Humans 0.04 (0–0.12) 0.48 (0.32–0.59)
LGDs (unsupervised) & Wolves 0.22 (0.13–0.30) 0.42 (0.27–0.58)
No evidence of wolf involvement in a human fatality in Greece 129
Predator criteria and factors assessment
We applied the ndings from the camera trapping, activity pattern analysis, eld ob-
servations and review of ocial and witness reports to the criteria and factors proposed
in Table 1 for the identication of the most likely responsible predator for the attack
(Table 4). From the 13 criteria/factors presented, sucient data for assessing them
were available for eleven. For two, the required data were either not analysed (i.e. canid
DNA collected from the attack site), not appropriately collected (i.e. DNA from the
mouths of suspected dogs) or were misinterpreted (i.e. the post-consumption patterns
of the victim’s corpse). We do not include in this assessment feral dogs as they were not
observed at the “Attack area”.
Figure 8. Activity overlap at the “Attack area” of humans (“recreationists” – i.e. hikers, bikers, hunters,
excluding researchers and shepherds) and (a) wolves (coecient of overlap Δ = 0.04) and (b) unsupervised
livestock guarding dogs (Δ = 0.55). Sunrise and sunset are marked at their average time for the study
period (GMT+2).
Yorgos Iliopoulos et al. / Nature Conservation 50: 115–143 (2022)
130
Table 4. Assessment of study ndings according to the proposed criteria and factors to the case for the
identication of either LGDs or wolves as responsible for the fatal attack considering the study ndings
(symbols: check () denotes evidence of a factor/criterion pointing at the species’ involvement, (X) to lack
of evidence, dash (–) not applicable in the current case study and (?) to inconclusive evidence).
1. Encounter rate between carnivore and human
Livestock Guarding Dogs Wolves
Facts Facts
• LGDs were active around the clock at “Attack area” • Wolves were detected only at night-time at “Attack area”
• e activity overlaps of unsupervised LGDs and humans (hikers,
hunters, cyclists) was high
• e activity overlap of wolves and humans (hikers, hunters,
cyclists) was 14 times lower than the corresponding overlap of
LGDs.
• At the time of the attack, the activity overlap was at its peak • At the time of the attack, the activity overlap was practically zero
Assessment Assessment
Probability of LGDs encountering the victim during the time and location
of the attack was very high. Criterion is met.
Probability of wolves encountering the victim during the time and
location of the attack was very low. e observed dierences in the
activity pattern of the wolf pack in the two sites (i.e. strictly nocturnal
at the attack site, cathemeral near the pack’s rendezvous site) is
consistent with wolf avoidance of LGDs and human activity, both of
which were 6–7 times lower in “Broader area”. Criterion is not met.
X
2. Location of attack site in relation to carnivore territory
Livestock Guarding Dogs Wolves
Facts Facts
• e attack site is adjacent to the corral of the goat herd guarded by the
LGDs. e herd and, therefore, the LGDs were present at the day of the
attack and used daily the area throughout our study period
• e attack site is located, at a distance > 5 km from its rendezvous
site/home-site. e wolf pups appear at the “Attack area” only after
the pack entered the nomadic phase (Mills et al. 2008) in November,
which coincided with the period that the LGDs and the goat herd
left the seasonal corral.
• LGDs demonstrated increased territorial defence behaviour and aggression
towards non-familiar humans (Witness report + authors’ observations)
Assessment Assessment
LGDs were likely to have reacted towards the victim’s presence as an
intruder present at the core of their territory. Risk factor applies.
Since the attack location is not a critical area for the protection of
the ospring, it is unlikely that wolves would have reacted with
aggression for defensive purposes. Risk factor does not apply
X
3. Carnivore health condition
Livestock Guarding Dogs Wolves
Facts Facts
• Animals appear healthy and in good physical condition – i.e. no visibly
malnourished individuals
• Animals appear healthy and in good physical condition – i.e. no
visibly malnourished individuals
• No records of rabies in Greece since 2014 • No cases of rabid wolves in Greece during the 2012–2014 rabies
outbreak
• No records of rabies in Greece since 2014
Assessment Assessment
e condition of the LGDs observed does not suggest that impaired
hunting ability due to physical disability or disease (rabies) would have
acted as driver of the attack. Risk factor does not apply.
e condition of the wolves does not suggest that impaired hunting
ability due to physical disability or disease (rabies) would have acted
as driver of the attack. Risk factor does not apply.
X X
4. Canid group size
Livestock Guarding Dogs Wolves
Facts Facts
• Pack size of eleven LGDs • Pack size of two adult wolves until 11 Nov.
Assessment Assessment
e LGD pack was able to subdue a human. e pack had many adult
animals and, therefore, it is possible that just one of them initiated the
attack triggering the rest to escalate it eventually to a fatality. Risk factor
applies in analogy to dog pack size.
Wolves were able to physically subdue an adult human. Risk factor
applies.
No evidence of wolf involvement in a human fatality in Greece 131
5. Dog(s) body size
Livestock Guarding Dogs Wolves
Facts
Not applicable
• Most LGDs were large-bodied
Assessment
e presence of large-bodied animals in the LGD pack highly increased
risk of fatality in the event of an attack is. Risk factor applies.
–
6. Number of male individuals
Livestock Guarding Dogs Wolves
Facts Facts
• At least eight of the eleven LGDs were male. • One of the two adult wolves was male.
Assessment Assessment
e large number of male dogs in the pack suggests that high levels of
intra-group competition may have been present, which in turn is a factor
for increased risk of attacks to humans. We were unable to determine
whether the male dogs were neutered, but, in general, this is a rare practice
for LGDs in Greece. Risk factor applies.
is risk factor refers to LGDs mostly. However, it is worth noting
that at this time of the year, adult male wolves have low levels of
testosterone as it is not the mating season. erefore, they probably
would not demonstrate levels of interspecies aggression. Risk factor
does not apply or is not relevant
–
7. Wolf prey availability and wolf habituation
Livestock Guarding Dogs Wolves
Not applicable
Facts
• e natural prey availability (wild boar, roe deer) for wolves in the
study area is low.
• ere is high availability of anthropogenic food sources (livestock,
carrion, hunting dogs) in the area at the given season, which wolves
can both prey on and scavenge.
Assessment
e wolf pack was likely accustomed to anthropogenic food
sources and this could lead to human habituation. While the pack
is probably not food stressed, the overall food requirements for the
pack were high given the presence of the pups. Risk factor applies.
–
8. Season of attack related to wolf biological cycle
Livestock Guarding Dogs Wolves
Not applicable
Facts
Fatal attack happened in late September.
Assessment
Attack was outside the period from May–August where predatory
wolf attacks to humans in human dominated landscapes have been
more frequent. Risk factor does not apply.
– X
9. Dog socialisation to humans and previous aggression
Livestock Guarding Dogs Wolves
Facts
Not applicable
• Dog group in rural area, raised in a livestock fold. Minimum contact
with family members.
• Witnesses reported an LGD pack that matches the one at the attack site
in terms of size, location and timing, as having demonstrated aggressive
behaviour towards people passing by.
• Dog aggression demonstrated towards the authors.
Not applicable
• Extensive occasions where LGDs roamed unsupervised in the attack site
at all hours of the day.
Assessment
e above evidence suggests that, as is typical in Greece for LGDs, the pack
dogs have not been positively reinforced to human socialisation and that
the observed aggressive behaviour towards strangers is the norm and not
the exception. Risk factor applies.
–
Yorgos Iliopoulos et al. / Nature Conservation 50: 115–143 (2022)
132
10. Human supervision
Livestock Guarding Dogs Wolves
Facts
Not applicable
• LGDs roamed unsupervised in the attack site at all hours of the day.
• At the time of the attack, the probability of a human (hiker, hunter,
cyclist) encountering an unsupervised LGD of the pack was maximum.
Assessment
e probability of the victim encountering the herd’s LGDs at a time that
they were not eectively supervised by the shepherd, which could have
tried to interrupt the escalation of an attack, is high. Risk factor applies.
–
11. Bite patterns
Livestock guarding dogs Wolves
Facts Facts
• e near complete post-mortem consumption of the victim’s body means
that examination of bite mark patterns inicted during the attack was
impossible.
• e near complete post-mortem consumption of the victim’s body
means that examination of bite mark patterns inicted during the
attack was impossible.
Assessment Assessment
Not possible to assess. Not possible to assess.
– –
12. Post-mortem consumption patterns and rates
Facts
• Near complete consumption of victim’s body, disarticulation including decapitation, breaking of skull and large bones, scattering of body
remains over a large area.
• e above post-mortem consumption occurred within maximum 36 hours.
Livestock Guarding Dogs Wolves
• Body remains (fragment of skull) retrieved by police at the watering
trough of the goat herd, located 70 m from the attack site.
• Wolves had presence in the area during night-time only.
• e victim’s jacket and underwear were located 50 m from the corral. • Only two wolves recorded at the “Attack area” until 12 November
(1.5 months after the attack)
• Around the clock presence of LGDs at the attack site.
Assessment Assessment
Domestic dogs – including breeds smaller than the LGDs observed – are
capable of both killing a human and causing the type of extensive and
fast post-mortem damaged observed in the case (e.g. Borchelt et al. 1983;
Haglund et al. 1989; Fonseca and Palacios 2013; Fonseca et al. 2015).
While similar-sized dog breeds to wolves have approximately 25% less
powerful bite compared to wolves (Christiansen and Wroe 2007), even
medium-sized dogs can break the skull and large bones of a human (e.g.
Steadman and Worne 2007; Buschmann et al. 2011). Consumption of a
human corpse by dogs can commence immediately after death, if there is
trauma and blood (Rothschild and Schneider 1997). Notably, Avis (1999)
reported that, in one case, a dog pack killed in one attack two people and
consumed them immediately, even though they had been fed earlier by
their owner.
Wolves can cause the type of extensive and fast post-mortem
damage observed in this case. Since a) only two adult wolves were
recorded being active in the area and only at night, b) according
to literature, wild wolves which have not fed for many days can
consume at most 10 kg of biomass in one meal, following which
they need several hours for digestion before feeding again (Wilmers
and Stahler 2002; Peterson and Ciucci 2003), c) consumption
could be interrupted by the adjacent large LGD pack at the corral
and d) the rescue team searching for the victim was active in the
area as of the evening of the attack (including during the night),
it is questionable whether wolves alone could have achieved the
near complete post-mortem consumption observed in one meal.
However, translocation of body parts away from the attack site
to feed wolf pups cannot be excluded. Similarly, other scavengers
recorded at the attack site (jackals, foxes) could have scavenged
on the body. erefore, the observed post-mortem consumption
pattern and rate of the victim’s corpse cannot be used as a criterion
for suggesting wolf involvement in pre-mortem fatal attack.
erefore, the observed post-mortem consumption pattern and rate of the
victim’s corpse cannot be used as a criterion for excluding dog involvement
in pre-mortem fatal attack, as suggested by the coroner.
? ?
No evidence of wolf involvement in a human fatality in Greece 133
13. Sampling of genetic material from bites and wounds
Facts
• e police retrieved torn garments (jacket, trousers, shocks, shirt) of the victim with puncture wounds.
• Cheek swabs and hair samples were collected by a veterinarian from ten LGDs of the goat herd corralled next to the attack site after sedating
them, > 72 h after the attack.
• Animal hairs were retrieved on top of bone fragments found at the attack site.
• All genetic analysis conducted involved the extraction of human DNA from collected samples.
• No attempt to extract carnivore DNA (i.e. from animal hair, possible saliva from garment).
Livestock Guarding Dogs Wolves
Assessment Assessment
e epithelial DNA obtained from the cheeks of the LGDs using swabs
was at best collected no less than 72 h following the attack (pers. comm.).
e aim of this analysis was to determine whether the LGDs had bitten
or consumed the victim. e established way for using genetic analysis
to identify the culprit of an attack (carnivore on humans or livestock) is
to collect saliva from the puncture wounds on the victim (or in case of
humans, also on his/her clothes), (Sundqvist et al. 2007; Caniglia et al.
2013; Harms et al. 2015; Caniglia et al. 2016; López-Bao et al. 2017;
Fabbri et al. 2018; Plumer et al. 2018). is was unfortunately not done in
this case, even though the victim’s torn clothes were retrieved.
Since no saliva was obtained from the victim’s clothes, the genetic
analysis was unable to elucidate possible wolf involvement in the
attack. For example, in a fatal wolf attack case on a woman in Alaska
(Butler et al. 2011), genetic analysis of saliva obtained from both
body remains and clothes successfully identied not only the species,
but also the number of individuals involved in the attack.
Moreover, the time elapsed from the attack until the swab collection means
that any foreign genetic material could have deteriorated in the unstable
(warm, humid) condition of the animals’ mouths, especially if only biting
was involved. According to a recent study that tested exactly this point, it
was concluded that hexogen (i.e. victim’s) DNA could not be traced in the
mouths of dogs after 4 h (Iarussi et al. 2020). So, the absence of human
DNA detection in the swab analysis does not absolve the dogs as potential
predators or scavengers. Finally, even if human DNA had been detected
in the LGDs’ mouths, the interpretation of its source would have been
problematic – it could have been due to post-mortem consumption and
not necessarily the attack phase of the incident.
erefore, the protocol used was inappropriate and the criterion is
not informative.
erefore, the protocol used was inappropriate and the criterion is not
informative.
– –
Discussion
Based on the assessment of our ndings, there is substantial circumstantial evidence
that the fatal attack on the British tourist in Greece on 21 September 2017 could have
been caused by the livestock guarding dogs (LGDs) of the goat herd corralled adjacent
to the attack site. Our evidence includes high activity overlap between recreationists
and unsupervised LGDs (with it peaking at 5 pm, which is the same time as that of
the attack), lack of LGD supervision, large LGD pack size, high ratio of males among
LGDs, close vicinity of the attack site to the LGDs and the livestock corral, and a
record of LGD aggression towards humans as recently as a few days before the attack.
Specically, regarding LGD aggression, apart from the author’s own eld observations,
two Austrian tourists reported seeing a helmet-mounted video of two Belgian bikers
who were attacked by seven aggressive dogs on the same section of the trail a couple
days before the attack. is was reported as a written statement to WWF-Greece and
submitted to the police. It was also shared with us with their consent.
However, there are certain inherent limitations due to the nature of our study
which dictate that the LGDs’ involvement in the fatal attack is highly probable, but
Yorgos Iliopoulos et al. / Nature Conservation 50: 115–143 (2022)
134
not certain. is includes that our data collection unavoidably occurred after the at-
tack and, therefore, cannot with certainty reect the exact conditions on the day of the
attack. Additionally, the behaviour of the wolves could have changed through time,
due to the brief (72 h), but intense human activity near the attack site by search and
forensic teams. erefore, in principle, a wolf attack cannot be excluded despite the
very low probability supported by our ndings. In terms of posthumous body con-
sumption, our ndings suggest that the same dogs and/or wild scavengers, including
wolves, could have contributed.
Although our study conrmed the presence of golden jackals in the study area and
the coroner’s media statements and report suggested the species could be involved in
the fatal attack, we did not consider the species as a potential predator in our assess-
ment. We decided this because there is no literature reference of jackals fatally attack-
ing humans anywhere in the world and injury reports are limited to countries where
rabies is endemic and human and jackal densities are very high (i.e. India, Akhtar and
Chauhan 2009), whilst rabies in Greece is no longer endemic. Moreover, jackals were
exclusively nocturnal at the study area and exhibited no overlap with human activity.
Finally, the resident jackal family group consisted of just two adults (low density).
Our ndings stem from a set of criteria developed to assist, along with standard
forensic protocols, the predator identication in cases such as ours (i.e. nearly complete
body consumption and delayed location of remains). On the contrary, the coroner’s
conclusion was based primarily on the post-mortem consumption pattern observed
and the lack of human DNA detection in the mouths of the dogs. Importantly, there
was no dierentiation in the report between the two distinct phases of the incident
– the attack and the post-mortem consumption of the body. Recognising these two
phases is important, as the species responsible for the fatal attack may not consume
(or may not be the only one to consume) the body. erefore, in that particular case,
the post-mortem consumption pattern cannot be used as a criterion for excluding dog
responsibility for the fatal attack. Moreover, the reported form and extent of post-mor-
tem consumption can result from dog consumption according to forensic literature
(e.g. Borchelt et al. 1983; Haglund et al. 1989; Avis 1999; Steadman and Worne 2007;
Buschmann et al. 2011; Fonseca and Palacios 2012; Fonseca et al. 2015).
Regarding the search for human DNA in the dog cheek swabs, the results could
not be informative in this specic case. e sampling occurred > 72 h after the attack
and, critically, not from a location that would identify the attacker, especially if only
bites had been inicted on the victim. Several forensic publications recommend to
sample for predator DNA from the saliva left on the wounds and clothes of the victim,
soon after the attack (e.g. Sundqvist et al. 2007; Caniglia et al 2013; Harms et al. 2015;
Caniglia et al. 2016; López-Bao et al. 2017; Fabbri et al. 2018; Plumer et al. 2018). Re-
gardless, even if genetic material is collected using swabs from the mouth of suspected
animals, a recent study by Iarussi et al. (2020) showed that no traces of hexogen (i.e.
the victim’s) DNA could be traced in the swab samples after 4 h. erefore, the absence
of human DNA in the dog cheek swabs analysed in our case study is not informative
about the involvement or not of the dogs in the attack.
No evidence of wolf involvement in a human fatality in Greece 135
Results from this case study are in line with the main conclusion of Linnell et al.
(2021) that human fatalities from healthy wolves in Europe are extremely rare (e.g.
non-existent for the period 2015–2018), while rabies remains the most prevalent cause
for veried human injuries or deaths from wolves. Rabies is no longer endemic in
Greece, at least since 2015 (EFSA 2021), while not a single case of a rabid wolf has
been recorded in the country since rabies was rst detected in 2012. On the contrary,
in 2016 alone, at least 45 human fatalities caused by dogs were recorded in Europe
(Sarenbo and Svensson 2020).
We recognise that the rarity of wild carnivore fatal attacks on humans in Europe,
especially with such extensive posthumous consumption, complicated eorts to resolve
this case. Forensic science literature of similar cases recommends a more interdiscipli-
nary approach to the evaluation of the evidence, which includes also ecological and
environmental characteristics of the attack (e.g. Fonseca and Palacios 2012). e value
of such a broader approach to the case was recognised by the Police Department, Forest
Service and Ministry of the Environment, which expeditiously provided the necessary
permits and support for our study. Moreover, the Public Prosecutor’s oce took into
consideration an earlier version of this report, amongst all other information available,
prior to bringing the owner of the goat herd to court with the accusation of involun-
tary manslaughter. After many adjournments of the court, the case verdict was issued
on September 23, 2022. e judges concluded that the death was due to attack by the
livestock guarding dogs of the goat herd being kept at the time at the temporary coral
near the attack site, and found the owner guilty of involuntary manslaughter.
Resolving the case is important for carnivore conservation in Europe, as it received
considerable national and international media coverage (Arbieu et al. 2021). e al-
leged involvement of wolves, despite the lack of concrete evidence to support it, has
left a lingering impression to the public regarding the threat that wolves pose to hu-
mans, as deduced by the frequent enquiries addressed to the authors regarding the case.
ough rare, even alleged wildlife involvement in human attacks have the potential to
decrease public tolerance towards these species, which can have broader repercussions
on wildlife conservation (Bombieri et al. 2018). erefore, we consider the detailed
description of what transpired in the death of the British tourist to be of value to the
broader public, forensic authorities and scientic/wildlife management community.
Our assessment regarding the responsibility of LGDs in the fatal attack is also not
without potential negative impacts on large carnivore conservation. LGDs are recog-
nised as an eective and socially acceptable tool for wildlife-human co-existence (Rigg
et al. 2011). e case could amplify existing concerns about the broader risk that
LGDs may pose to recreationists (Linnell and Lescureux 2015). erefore, ensuring
both adequate livestock protection from large carnivores and public safety is important
in areas where nature tourism and extensive livestock raising co-occur, since both ac-
tivities are important for rural economies. In Greece, a large LGD group size, like the
one observed in this case, is common in areas with large carnivores (wolf and brown
bear). Moreover, LGDs often need to work independently from the shepherd’s supervi-
sion to locate and deter predators (Landry et al. 2020).
Yorgos Iliopoulos et al. / Nature Conservation 50: 115–143 (2022)
136
e location of the seasonal livestock corral next to a tourist trail was a decisive fac-
tor in the case, as it greatly increased the chance of the LGDs encountering the victim.
In addition, the victim’s response to approaching dogs, though unknown, could have
contributed to the initiation and escalation of the attack, as documented in similar
studies (e.g. Borchelt et al. 1983; Rezac et al. 2015; Reese and Vertalka 2020).
Conclusions
Based on the assessment of our ndings, we conclude that there is no concrete evidence
that wolves were responsible for the death of the British tourist in the Maroneia-Petrota
region, northern Greece. On the contrary, evidence provided by the assessable risk crite-
ria, based on literature review and eld observations, suggest an increased probability of
a dog attack as a probable cause of death of the British tourist. e court’s decision issued
around the same time as the acceptance of this article was also in line with our assessment.
ere are lessons to be learned from this tragic case study. Risks related to livestock
guarding dogs need to be carefully evaluated in areas with frequent tourist activity to
avoid human injuries and fatalities. We propose three practical measures to reduce
the probability of future LGD attacks on humans. First, all permanent and seasonal
livestock corrals in the vicinity of tourist areas (e.g. hiking trails, open-air archaeologi-
cal sites, mountain lodges) should be mapped, their potential risk to users evaluated
including assessment of dog behaviour and, if necessary, relocated. In all cases, warning
signs can be added both on site and in hiking maps or applications. Secondly, together
with the guidelines provided for encounters with potentially dangerous wildlife (e.g.
brown bears; Bombieri et al. 2019), the public should also be informed about best
ways of avoiding and handling dangerous LGD encounters. Additionally, in countries
where deterrence tools, such as pepper sprays, are not allowed, legal exceptions to their
use for animal deterrence in rural areas could be considered.
Finally, a multidisciplinary approach in the assessment of putative wildlife attacks
on humans can reduce misidentications of the responsible species and, therefore, pre-
vent unfounded decrease in public tolerance for large carnivores, which is dicult to
achieve and even harder to maintain. To reduce future controversies, police routines
should be established that automatically integrate wildlife expertise into such investiga-
tions, as well as establishing best practices for the collection of appropriate forensic data.
Acknowledgements
e study was implemented under a research licence from the Hellenic Ministry of En-
vironment (no. 154593/2487). e authors would like to thank the Head of the Rho-
dope Forest Directorate, M. Gotzaridou for assistance and support during eld work,
Sapes Police Department for facilitating ongoing research, P. Maragou from WWF
Greece for providing access to testimonials from the study area, R.A. Montgomery and
No evidence of wolf involvement in a human fatality in Greece 137
three other anonymous reviewers for providing valuable feedback on an earlier version
of this manuscript and Callisto Wildlife Society for providing all necessary equipment
needed for the research.
References
Akhtar N, Chauhan NPS (2009) Food habitats and human-jackal interaction in Marwahi For-
est Division, Bilaspur Chhattisgarh. Indian Forester (October): 1347–1356. https://www.
cabdirect.org/globalhealth/abstract/20103013807
Ambarli H (2019) Analysis of wolf–human conicts: Implications for damage mitigation meas-
ures. European Journal of Wildlife Research 65(6): 1–9. https://doi.org/10.1007/s10344-
019-1320-4
Arbieu U, Chapron G, Astaras C, Bunnefeld N, Harkins S, Iliopoulos Y, Mehring M, Re-
inhardt I, Mueller T (2021) News selection and framing: e media as a stakeholder in
human-carnivore coexistence. Environmental Research Letters 16(6): 064075. https://doi.
org/10.1088/1748-9326/ac05ef
Avis SP (1999) Dog pack attack: Hunting humans. e American Journal of Forensic Medicine
and Pathology 20(3): 243–246. https://doi.org/10.1097/00000433-199909000-00005
Behdarvand N, Kaboli M (2015) Characteristics of Gray Wolf Attacks on Humans in an Al-
tered Landscape in the West of Iran. Human Dimensions of Wildlife 20(2): 112–122.
https://doi.org/10.1080/10871209.2015.963747
Behdarvand N, Kaboli M, Ahmadi M, Nourani E, Salman Mahini A, Asadi Aghbolaghi M
(2014) Spatial risk model and mitigation implications for wolf–human conict in a highly
modied agroecosystem in western Iran. Biological Conservation 177: 156–164. https://
doi.org/10.1016/j.biocon.2014.06.024
Boitani L (2018) Canis lupus (errata version published in 2019). e IUCN Red List of reat-
ened Species 2018: e.T3746A144226239. [Accessed on 08 February 2022]
Bombieri G, Nanni V, Delgado M del M, Fedriani JM, López-Bao JV, Pedrini P, Penteriani V
(2018) Content Analysis of Media Reports on Predator Attacks on Humans: Toward an
Understanding of Human Risk Perception and Predator Acceptance. Bioscience 68(8):
577–584. https://doi.org/10.1093/biosci/biy072
Bombieri G, Naves J, Penteriani V, Selva N, Fernández-Gil A, López-Bao JV, Ambarli H, Bau-
tista C, Bespalova T, Bobrov V, Bolshakov V, Bondarchuk S, Camarra JJ, Chiriac S, Ciucci
P, Dutsov A, Dykyy I, Fedriani JM, García-Rodríguez A, Garrote PJ, Gashev S, Gro C,
Gutleb B, Haring M, Härkönen S, Huber D, Kaboli M, Kalinkin Y, Karamanlidis AA,
Karpin V, Kastrikin V, Khlyap L, Khoetsky P, Kojola I, Kozlow Y, Korolev A, Korytin N,
Kozsheechkin V, Krofel M, Kurhinen J, Kuznetsova I, Larin E, Levykh A, Mamontov V,
Männil P, Melovski D, Mertzanis Y, Meydus A, Mohammadi A, Norberg H, Palazón S,
Pătrașcu LM, Pavlova K, Pedrini P, Quenette PY, Revilla E, Rigg R, Rozhkov Y, Russo
LF, Rykov A, Saburova L, Sahlén V, Saveljev AP, Seryodkin IV, Shelekhov A, Shishikin A,
Shkvyria M, Sidorovich V, Sopin V, Støen O, Stok J, Swenson JE, Tirski D, Vasin A, Wa-
bakken P, Yarushina L, Zwijacz-Kozica T, Delgado MM (2019) Brown bear attacks on hu-
Yorgos Iliopoulos et al. / Nature Conservation 50: 115–143 (2022)
138
mans: A worldwide perspective. Scientic Reports 9(1): e8753. https://doi.org/10.1038/
s41598-019-44341-w
Borchelt PL, Lockwood R, Beck AM, Voith VL (1983) Attacks by packs of dogs involving predation
on human beings. Public Health Reports 98: 57–66. https://www.jstor.org/stable/4627350
Boyce MS (2018) Wolves for Yellowstone: Dynamics in time and space. Journal of Mammalogy
99(5): 1021–1031. https://doi.org/10.1093/jmammal/gyy115
Buschmann C, Solarino B, Püschel K, Czubaiko F, Heinze S, Tsokos M (2011) Post-mortem
decapitation by domestic dogs: ree case reports and review of the literature. Forensic
Science, Medicine, and Pathology 7(4): 344–349. https://doi.org/10.1007/s12024-011-
9233-x
Butler L, Dale B, Beckmen K, Farley S (2011) Findings related to the March 2010 fatal wolf
attack near Chignik Lake, Alaska. Wildlife Special Publication, Alaska departement of sh
and game, Division of wildlife conservation, USA. https://www.adfg.alaska.gov/static/
home/news/pdfs/wolfattackfatality.pdf
Caniglia R, Fabbri E, Mastrogiuseppe L, Randi E (2013) Who is who? Identication of live-
stock predators using forensic genetic approaches. Forensic Science International. Genetics
7(3): 397–404. https://doi.org/10.1016/j.fsigen.2012.11.001
Caniglia R, Galaverni M, Delogu M, Fabbri E, Musto C, Randi E (2016) Big bad wolf or man’s
best friend? Unmasking a false wolf aggression on humans. Forensic Science International.
Genetics 24: e4–e6. https://doi.org/10.1016/j.fsigen.2016.06.009
Chapron G, Kaczensky P, Linnell JDC, von Arx M, Huber D, Andrén H, López-Bao JV, Ad-
amec M, Álvares F, Anders O, Balčiauskas L, Balys V, Bedő P, Bego F, Blanco JC, Breiten-
moser U, Brøseth H, Bufka L, Bunikyte R, Ciucci P, Dutsov A, Engleder T, Fuxjäger C,
Gro C, Holmala K, Hoxha B, Iliopoulos Y, Ionescu O, Jeremić J, Jerina K, Kluth G,
Knauer F, Kojola I, Kos I, Krofel M, Kubala J, Kunovac S, Kusak J, Kutal M, Liberg O,
Majić A, Männil P, Manz R, Marboutin E, Marucco F, Melovski D, Mersini K, Mertza-
nis Y, Mysłajek RW, Nowak S, Odden J, Ozolins J, Palomero G, Paunović M, Persson J,
Potočnik H, Quenette P-Y, Rauer G, Reinhardt I, Rigg R, Ryser A, Salvatori V, Skrbinšek
T, Stojanov A, Swenson JE, Szemethy L, Trajçe A, Tsingarska-Sedefcheva E, Váňa M,
Veeroja R, Wabakken P, Wöl M, Wöl S, Zimmermann F, Zlatanova D, Boitani L (2014)
Recovery of large carnivores in Europe’s modern human-dominated landscapes. Science
346(6216): 1517–1519. https://doi.org/10.1126/science.1257553
Christiansen P, Wroe S (2007) Bite forces and evolutionary adaptations to feeding ecology in car-
nivores. Ecology 88(2): 347–358. https://doi.org/10.1890/0012-9658(2007)88[347:BFA
EAT]2.0.CO;2
Cimatti M, Ranc N, Benítez-López A, Maiorano L, Boitani L, Cagnacci F, Čengić M, Ciucci
P, Huijbregts MAJ, Krofel M, López-Bao JV, Selva N, Andren H, Bautista C, Ćirović D,
Hemmingmoore H, Reinhardt I, Marenče M, Mertzanis Y, Pedrotti L, Trbojević I, Zetter-
berg A, Zwijacz-Kozica T, Santini L (2021) Large carnivore expansion in Europe is associ-
ated with human population density and land cover changes. Diversity & Distributions
27(4): 602–617. https://doi.org/10.1111/ddi.13219
de Bruxelles S, Carassava A (2017) Wolves blamed for death of mauled British tourist Celia
Hollingworth. e Times. https://www.thetimes.co.uk/article/wolves-blamed-for-british-
tourist-s-death-x708crgcp [Assesed February 8, 2021]
No evidence of wolf involvement in a human fatality in Greece 139
EFSA [European Food Safety Authority and European Centre for Disease Prevention and
Control] (2021) e European Union One Health 2019 Zoonoses Report. EFSA Journal
19(2): e06406 [286 pp]. https://doi.org/10.2903/j.efsa.2021.6406
“Express” (2017) Horror over British woman mauled to death by pack of wild dogs in Greece.
https://www.express.co.uk/news/world/858874/Dog-Attack-Greece-Celia-Hollingworth
[Accessed February 8, 2022]
Fabbri E, Velli E, D’Amico F, Galaverni M, Mastrogiuseppe L, Mattucci F, Caniglia R (2018)
From predation to management: monitoring wolf distribution and understanding depre-
dationpatterns from attacks on livestock. Hystrix, the Italian Journal of Mammalogy 29:
101–110. https://doi.org/10.4404/HYSTRIX-00070-2018
Fegan-Earl AW (2005) Animal attacks and injuries. Fatal and non-fatal. In: Byard R, Corey T,
Henderson C, Payne-James J (Eds) Encyclopedia of forensic and legal medicine, Vol. 1, 1st
edn. Oxford, UK, 68–71. https://doi.org/10.1016/B0-12-369399-3/00013-6
Fonseca GM, Palacios R (2012) An unusual case of predation: Dog pack or cougar attack? Journal
of Forensic Sciences 58(1): 224–227. https://doi.org/10.1111/j.1556-4029.2012.02281.x
Fonseca GM, Mora E, Lucena J, Cantin M (2015) Forensic studies of dog attacks on humans:
A focus on bite mark analysis. Research and Reports in Forensic Medical Science 5: 39–51.
https://doi.org/10.2147/RRFMS.S92068
Gholami A, Fayaz A, Farahtaj F (2014) Rabies in Iran: Past, Present and Future. Journal of
Medical Microbiology and Infectious Diseases 2(1): 1–10. http://jommid.pasteur.ac.ir/
article-1-49-en.html
Haglund WD, Reay DT, Swindler DR (1989) Canid Scavenging/Disarticulation Sequence of
Human Remains in the Pacic Northwest. Journal of Forensic Sciences 34(3): e12679J.
https://doi.org/10.1520/JFS12679J
Haidt A, Gawryś R, Szewczyk M (2021) Human Decision-Making as a Key Factor in the Risk
of Wolf–Dog Interactions during Outdoor Activities. Animals 11(9): 2497. https://doi.
org/10.3390/ani11092497
Harms V, Nowak C, Carl S, Muñoz-Fuentes V (2015) Experimental evaluation of genetic
predator identication from saliva traces on wildlife kills. Journal of Mammalogy 96(1):
138–143. https://doi.org/10.1093/jmammal/gyu014
Heilhecker E, iel RP, Wayne H (2007) Wolf, Canis lupus, behavior in areas of frequent human ac-
tivity. Canadian Field Naturalist 121(3): 256–260. https://doi.org/10.22621/cfn.v121i3.472
Heine H (2017) Streit um Wölfe: Landwirtschaftsminister will Abschuss erleichtern. Der Ta-
gesspiegel. https://www.tagesspiegel.de/berlin/streit-um-woelfe-landwirtschaftsminister-
will-abschuss-erleichtern/20400392.html [Accessed February 8, 2022]
Homann CF, Montgomery RA, Jepson PR (2017) Examining the eect of billboards in shap-
ing the great wolf debate of the American West. Human Dimensions of Wildlife 22(3):
267–281. https://doi.org/10.1080/10871209.2017.1314059
Hsu Y, Sun L (2010) Factors associated with aggressive responses in pet dogs. Applied Animal
Behaviour Science 123(3–4): 108–123. https://doi.org/10.1016/j.applanim.2010.01.013
Iarussi F, Cipolloni L, Bertozzi G, Sasso L, Ferrara M, Salerno M, Rubino GTR, Maglietta F,
Dinisi A, Albano D, Iarussi V, Pomara C, Sessa F (2020) Dog-bite-related attacks: A new
forensic approach. Forensic Science International 310: 110254. https://doi.org/10.1016/j.
forsciint.2020.110254
Yorgos Iliopoulos et al. / Nature Conservation 50: 115–143 (2022)
140
Iliopoulos Y, Youlatos D, Sgardelis S (2014) Wolf pack rendezvous site selection in Greece is
mainly aected by anthropogenic landscape features. European Journal of Wildlife Re-
search 60(1): 23–34. https://doi.org/10.1007/s10344-013-0746-3
Isloor S, Marissen WM, Veeresh BH, Nithinprabhu K, Kuzmin IV, Rupprecht CE, Satyanaray-
ana ML, Deepti BR, Sharada R, Neelufer MS, Yathiraj S, Abdul Rahman S (2014) First
Case Report of Rabies in a Wolf (Canis lupus pallipes) from India. Journal of Veterinary
Medical Research 001: 1–6.
Janeiro-Otero A, Newsome TM, Van Eeden LM, Ripple WJ, Dormann CF (2020) Grey wolf
(Canis lupus) predation on livestock in relation to prey availability. Biological Conservation
243: 108433. https://doi.org/10.1016/j.biocon.2020.108433
Jhala YV, Sharma DK (1997) Child-lifting by wolves in eastern Uttar Pradesh, India. Journal
of Wildlife Research 2: 94–101. https://ci.nii.ac.jp/naid/10026778633/ [Accessed August
7, 2021]
Kansky R, Knight AT (2014) Key factors driving attitudes towards large mammals in con-
ict with humans. Biological Conservation 179: 93–105. https://doi.org/10.1016/j.bio-
con.2014.09.008
Kneafsey B, Condon KC (1995) Severe dog-bite injuries, introducing the concept of pack
attack: A literature review and seven case reports. Injury 26(1): 37–41. https://doi.
org/10.1016/0020-1383(95)90550-H
Kuijper DPJ, Churski M, Trouwborst A, Heurich M, Smit C, Kerley GIH, Cromsigt JPGM (2019)
Keep the wolf from the door: How to conserve wolves in Europe’s human-dominated land-
scapes? Biological Conservation 235: 102–111. https://doi.org/10.1016/j.biocon.2019.04.004
Landry JM, Borelli JL, Drouilly M (2020) Interactions between livestock guarding dogs and
wolves in the southern French Alps. Journal of Vertebrate Biology 69(3): 20078 [1–18].
https://doi.org/10.25225/jvb.20078
Lescureux N, Linnell JDC (2014) Review. Biological Conservation Complete 171: 232–245.
https://doi.org/10.1016/j.biocon.2014.01.032
Linnell JDC, Alleau J (2015) Predators that kill humans: Myth, reality, context and the politics
of wolf attacks on people. Problematic Wildlife: A Cross-Disciplinary Approach, 357–371.
https://doi.org/10.1007/978-3-319-22246-2_17
Linnell JDC, Lescureux N (2015) Livestock guarding dogs – cultural heritage icons with a new
relevance for mitigating conservation conicts. Norwegian Institute for Nature Research,
Trondheim.
Linnell JDC, Andersen R, Andersone Z, Balciauskas L, Blanco JC, Boitani L, Brainerd S, Be-
itenmoser U, Kojola I, Liberg O, Løe J, Okarma H, Pedersen HC, Promberger C, Sand H,
Solberg EJ, Valdmann H, Wabakken P (2002) e fear of wolves: A review of wolfs attacks
on humans. NINA Oppdragsmelding 731, Norsk institutt for naturforskning NINA.
Linnell JDC, Solberg EJ, Brainerd S, Liberg O, Sand H, Wabakken P, Kojola I (2003) Is the
Fear of Wolves Justied? A Fennoscandian Perspective. Acta Zoologica Lituanica 13(1):
34–40. https://doi.org/10.1080/13921657.2003.10512541
Linnell JDC, Kovtun E, Rouart I (2021) Wolf attacks on humans: an update for 2002–2020.
NINA Report 1944, Norwegian Institute for Nature Research.
Löe J, Röskaft E (2004) Large carnivores and human safety: A review. Ambio 33(6): 283–288.
https://doi.org/10.1579/0044-7447-33.6.283
No evidence of wolf involvement in a human fatality in Greece 141
López-Bao JV, Frank J, Svensson L, Åkesson M, Langefors Å (2017) Building public trust in
compensation programs through accuracy assessments of damage verication protocols.
Biological Conservation 213: 36–41. https://doi.org/10.1016/j.biocon.2017.06.033
Marion M, Béata C, Sarcey G, Delfante S, Marlois N (2018) Study of aggressiveness in live-
stock-guarding dogs based on rearing method. Journal of Veterinary Behavior 25: 14–16.
https://doi.org/10.1016/j.jveb.2018.03.001
Martin HW, Mech LD, Fieberg J, Metz MC, Macnulty DR, Stahler DR, Smith DW (2018)
Factors aecting gray wolf (Canis lupus) encounter rate with elk (Cervus elaphus) in Yel-
lowstone National Park. Canadian Journal of Zoology 96(9): 1032–1042. https://doi.
org/10.1139/cjz-2017-0220
Matos RE, Jakuba T, Mino I, Fejsakova M, Demeova A, Kottferova J (2015) Characteristics
and risk factors of dog aggression in the Slovak Republic. Veterinární Medicína 60(8):
432–445. https://doi.org/10.17221/8418-VETMED
McNay ME (2002) Wolf-Human Interactions in Alaska and Canada: A Review of the Case
History. Wildlife Society Bulletin 30: 831–843. https://www.jstor.org/stable/3784237
McNay ME (2007) A Review of Evidence and Findings Related to the Death of Kenton Car-
negie on 8 November 2005 Near Points North, Saskatchewan. Alaska Department of Fish
and Game, Fairbanks, Alaska.
McNay ME, Mooney PW (2005) Attempted Predation of a Child by a Gray Wolf, near Icy Bay,
Alaska. Canadian Field Naturalist 119(2): 197–201. https://doi.org/10.22621/cfn.v119i2.106
Meredith M, Ridout M (2021) Overview of the overlap package: https://cran.r-project.org/
web/packages/overlap/vignettes/overlap.pdf [Accessed August 8, 2021]
Mikkola S, Salonen M, Puurunen J, Hakanen E, Sulkama S, Araujo C, Lohi H (2021) Ag-
gressive behaviour is aected by demographic, environmental and behavioural factors in
purebred dogs. Scientic Reports 11: 1–10. https://doi.org/10.1038/s41598-021-88793-5
Mills KJ, Patterson BR, Murray DL (2008) Direct Estimation of Early Survival and Movements
in Eastern Wolf Pups. e Journal of Wildlife Management 72(4): 949–954. https://doi.
org/10.2193/2006-457
Muhly TB, Semeniuk C, Massolo A, Hickman L, Musiani M (2011) Human activity helps
prey win the predator-prey space race. PLoS ONE 6(3): e17050. https://doi.org/10.1371/
journal.pone.0017050
Notari L, Cannas S, Di Sotto YA, Palestrini C (2020) A retrospective analysis of dog–dog
and dog–human cases of aggression in Northern Italy. Animals 10(9): 1–17. https://doi.
org/10.3390/ani10091662
Nowak S, Szewczyk M, Tomczak P, Całus I, Figura M, Mysłajek RW (2021) Social and envi-
ronmental factors inuencing contemporary cases of wolf aggression towards people in Po-
land. European Journal of Wildlife Research 67(4): 69. https://doi.org/10.1007/s10344-
020-01455-1
Patronek GJ, Sacks JJ, Delise KM, Cleary DV, Marder AR (2013) Co-occurrence of potentially
preventable factors in 256 dog bite-related fatalities in the United States (2000–2009).
Journal of the American Veterinary Medical Association 243(12): 1726–1736. https://doi.
org/10.2460/javma.243.12.1726
Penteriani V, Delgado M del M, Pinchera F, Naves J, Fernández-Gil A, Kojola I, Härkönen
S, Norberg H, Frank J, Fedriani JM, Sahlén V, Støen O-G, Swenson JE, Wabakken P,
Yorgos Iliopoulos et al. / Nature Conservation 50: 115–143 (2022)
142
Pellegrini M, Herrero S, López-Bao JV (2016) Human behaviour can trigger large carni-
vore attacks in developed countries. Scientic Reports 6: 1–8. https://doi.org/10.1038/
srep20552
Penteriani V, Bombieri G, Fedriani JM, López-Bao JV, Garrote PJ, Russo LF, Delgado M del
M (2017) Humans as Prey: Coping with Large Carnivore Attacks using a Predator-Prey
Interaction Perspective. Human-Wildlife Interactions 11: 10. https://doi.org/https://doi.
org/10.26077/dpyb-q611
Peterson RO, Ciucci P (2003) e wolf as a carnivore. In: Mech LD, Boitani L (Eds) Wolves: be-
haviour, ecology and conservation. e University of Chicago Press, Chicago, USA, 161–191.
Plumer L, Talvi T, Männil P, Saarma U (2018) Assessing the roles of wolves and dogs in livestock
predation with suggestions for mitigating human–wildlife conict and conservation of wolves.
Conservation Genetics 19(3): 665–672. https://doi.org/10.1007/s10592-017-1045-4
R Core Team (2019). A language and environment for statistical computing. R Foundation for
Statistical Computing, Vienna.
Raghavan M (2008) Fatal dog attacks in Canada, 1990–2007. e Canadian Veterinary Jour-
nal 49: 577. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2387261/ [Accessed August
8, 2021]
Rajpurohit KS (1999) Child lifting: Wolves in Hazaribagh, India. Ambio 28: 162–166.
Reese LA, Vertalka JJ (2020) Understanding Dog Bites: e Important Role of Human Behav-
ior. Journal of Applied Animal Welfare Science 24(4): 331–346. https://doi.org/10.1080/
10888705.2020.1790371
Rezac P, Rezac K, Slama P (2015) Human behavior preceding dog bites to the face. Veterinary
Journal 206(3): 284–288. https://doi.org/10.1016/j.tvjl.2015.10.021
Ridout MS, Linkie M (2009) Estimating overlap of daily activity patterns from camera trap
data. Journal of Agricultural Biological & Environmental Statistics 14(3): 322–337.
https://doi.org/10.1198/jabes.2009.08038
Rigg R, Finďo S, Wechselberger M, Gorman ML, Sillero-Zubiri C, Macdonald DW (2011)
Mitigating carnivore–livestock conict in Europe: Lessons from Slovakia. Oryx 45(2):
272–280. https://doi.org/10.1017/S0030605310000074
Ripple WJ, Beschta RL (2012) Trophic cascades in Yellowstone: e rst 15 years after wolf
reintroduction. Biological Conservation 145(1): 205–213. https://doi.org/10.1016/j.bio-
con.2011.11.005
Roll A, Unshelm J (1997) Aggressive conicts amongst dogs and factors aecting them.
Applied Animal Behaviour Science 52(3–4): 229–242. https://doi.org/10.1016/S0168-
1591(96)01125-2
Rothschild MA, Schneider V (1997) On the temporal onset of postmortem animal scavenging
“Motivation” of the animal. Forensic Science International 89(1–2): 57–64. https://doi.
org/10.1016/S0379-0738(97)00112-6
Rubin HD, Beck AM (1982) Ecological behavior of free-ranging urban pet dogs. Applied Ani-
mal Ethology 8(1–2): 161–168. https://doi.org/10.1016/0304-3762(82)90141-9
Santoro V, Smaldone G, Lozito P, Smaldone M, Introna F (2011) A forensic approach to
fatal dog attacks. A case study and review of the literature. Forensic Science International
206(1–3): e37–e42. https://doi.org/10.1016/j.forsciint.2010.07.026
No evidence of wolf involvement in a human fatality in Greece 143
Sarenbo S, Svensson PA (2020) Bitten or struck by dog: A rising number of fatalities in Eu-
rope, 1995–2016. Forensic Science International 18: 110592. https://doi.org/10.1016/j.
forsciint.2020.110592
Shields LBE, Bernstein ML, Hunsaker JC, Stewart DM (2009) Dog bite-related fatalities: A 15-
year review of Kentucky medical examiner cases. e American Journal of Forensic Medi-
cine and Pathology 30(3): 223–230. https://doi.org/10.1097/PAF.0b013e3181a5e558
Shuler CM, DeBess EE, Lapidus JA, Hedberg K (2008) Canine and human factors related to
dog bite injuries. Journal of the American Veterinary Medical Association 232(4): 542–
546. https://doi.org/10.2460/javma.232.4.542
Sidorov GN, Sidorova DG, Poleshchuk EM (2010) Rabies of wild mammals in Russia at the
turn of the 20th and 21st centuries. Biology Bulletin of the Russian Academy of Sciences
37(7): 684–694. https://doi.org/10.1134/S1062359010070034
Sidorovich VE, Tikhomirova LL, Jȩdrzejewska B (2003) Wolf Canis lupus numbers, diet
and damage to livestock in relation to hunting and ungulate abundance in northeastern
Belarus during 1990–2000. Wildlife Biology 9(2): 103–111. https://doi.org/10.2981/
wlb.2003.032
Steadman DW, Worne H (2007) Canine scavenging of human remains in an indoor set-
ting. Forensic Science International 173(1): 78–82. https://doi.org/10.1016/j.forsci-
int.2006.11.011
Sundqvist A-K, Ellegren H, Vilà C (2007) Wolf or dog? Genetic identication of predators
from saliva collected around bite wounds on prey. Conservation Genetics 9: 1275–1279.
https://doi.org/10.1007/s10592-007-9454-4
“e Guardian” (2017) British woman mauled to death by wild dogs in Greece is named.
https://www.theguardian.com/uk-news/2017/sep/25/british-woman-mauled-death-wild-
dogs-greece-named [Accessed February 8, 2022]
Turkmen S, Sahin A, Gunaydn M, Tatli O, Karaca Y, Turedi S, Gunduz A (2012) A Wild Wolf
Attack and Its Unfortunate Outcome: Rabies and Death. Wilderness & Environmental
Medicine 23(3): 248–250. https://doi.org/10.1016/j.wem.2012.03.009
Wang L, Tang Q, Liang G (2014) Rabies and rabies virus in wildlife in mainland China, 1990–
2013. International Journal of Infectious Diseases 25: 122–129. https://doi.org/10.1016/j.
ijid.2014.04.016
Wilmers CC, Stahler DR (2002) Constraints on active-consumption rates in gray wolves,
coyotes, and grizzly bears. Canadian Journal of Zoology 80(7): 1256–1261. https://doi.
org/10.1139/z02-112
Content uploaded by Yorgos Iliopoulos
Author content
All content in this area was uploaded by Yorgos Iliopoulos on Oct 20, 2022
Content may be subject to copyright.