Received: 20 August 2020

Revised: 11 December 2020

Accepted: 13 January 2021

DOI: 10.1111/1440-1703.12211

NOTES AND COMMENTS

Scavenging and carcass caching behavior by European
wildcat (Felis silvestris)
Miha Krofel1|

Damjan Južnič1 | Maximilian L. Allen2

1

Biotechnical Faculty, Department ofForestry, University of Ljubljana,
Ljubljana, Slovenia

2

Abstract
While scavenging has been repeatedly reported for several felid species, sur-Illinois Natural History Survey,
University of Illinois, Champaign, Illinois

prisingly little information is available on scavenging behavior of theEuropean wildcat (Felis silvestris). To fill this knowledge gap, we used camera

Correspondence
Miha Krofel, University of Ljubljana,
Biotechnical Faculty, Department ofForestry, Ljubljana, Slovenia.
Email: miha.krofel@bf.uni-lj.si; miha.
krofel@gmail.com

traps to document scavenging behavior at the 48 experimentally set deer carcasses at random locations throughout the year. We recorded European wild-

Funding information
Javna Agencija za Raziskovalno Dejavnost
RS, Grant/Award Numbers: N1-0163,
P4-0059

cats scavenging on 38% of the eight carcasses set in winter and on none set in
the other parts of the year. Wildcats fed on two carcasses for extended periods(up to 22 days) with an average of 3.3 visits per day and 7.8-h interval between
the visits. We recorded scavenging throughout the day, but analysis indicateda crepuscular pattern. We also recorded caching behavior on 7% of the visits(n = 105), when wildcats used leaves or snow to partly or completely cover the
carcasses. Beside wildcats, 12 other vertebrate species of scavengers wererecorded at the carcasses. We recorded agonistic interaction with European
badger (Meles meles) and despite its smaller size, the wildcat managed todefend the carcass. The extensive feeding, frequent caching behavior andactive defense from scavengers indicate that the wildcats recognized the ungulate carcasses as an important food source in winter and that scavenging could
be a neglected aspect of the European wildcat ecology. We also suggest thatcaching behavior could be regularly used by the European wildcat when feeding on larger carcasses, but was likely previously missed due to limited
research effort to record scavenging and caching behavior.
KEYWORDS
camera traps, felids, food caching, interference competition, scavengers

1 | INTRODUCTION
Scavenging, where organisms from microbes to vertebrates
compete to consume ephemeral carrion, is an important
ecological process (Sebastián-González et al., 2019;
Wilmers, Stahler, Crabtree, Smith, & Getz, 2003). Scavengers provide functions in ecosystems and to humanity byremoving decaying carrion and potentially harmful pathoc, Penezic, &gens, as well as stabilizing food webs (Cirovi
556© 2021 The Ecological Society of Japan

Krofel, 2016; Sebastián-González et al., 2020; Wilson &Wolkovich, 2011). At the same time, carrion is an inexpensive resource that is used as a nutritional supplement in
the diet of many facultative scavengers (Allen, Elbroch,
Wilmers, & Wittmer, 2015; Inagaki et al., 2020), especiallyduring winter when other food sources are less available(Krofel, 2011; Selva, Jedrzejewska, Jedrzejewski, &Wajrak, 2005; Wilmers et al., 2003). Despite the benefits offeeding on carrion, there are also risks of injury or death,wileyonlinelibrary.com/journal/ere

Ecological Research. 2021;36:556–561.

KROFEL ET AL.

since fresh carcasses often attract several scavengers,which results in increased probability for agonistic
inter- or intra-specific interactions (Allen, Wilmers,
Elbroch, Golla, & Wittmer, 2016). Many scavengers andpredators have developed various behavioral adjustments to reduce the competition, such as caching thecarcass in trees, in caves or covering it with various
materials (Balme, Miller, Pitman, & Hunter, 2017;
Bischoff-Mattson & Mattson, 2009; de Ruiter &Berger, 2001; Krofel, Skrbinšek, & Mohorovic, 2019).
Felids are mostly hyper-carnivorous specialized
hunters with dentition and other morphological adaptations specialized for hunting (Van Valkenburgh, 1989).
However, many felid species readily scavenge when carcasses are available (Hunter, 2015). The proportion offood obtained through scavenging is generally considered
to be lower compared to the more generalist carnivores(e.g., hyaenids, ursids, canids and mustelids), but can still
be substantial for some felids. Scavenging is frequently
reported for the larger felids of the Panthera lineage,
while much less is known about this behavior among thesmaller cats. Scavenging has been reported for several of
the smaller felid species (Hunter, 2015), including theAfrican wildcat (Felis lybica) (Herbst, 2009), sand cat(F. margarita) (Brighten & Burnside, 2019) and blackfooted cat (Felis nigripes) (Renard, Lavoie, Pitt, &Larivière, 2015; Sliwa, 1994), many of which were also
reported caching larger kills or carcasses. It remains
unknown whether the lower frequency of reporting isconnected with lower frequency of scavenging amongsmaller felids or is this due to a lack of studies on scavenging behavior of the smaller felids.
Surprisingly little information is available on scavenging behavior of the European wildcat (Felis silvestris).
European wildcats occur predominantly in forested landscapes of Europe and feed mostly on rodents and othersmall prey (Hunter, 2015). While about half of the wildcat dietary studies in Europe reported consumption of
ungulates, they typically represent less than 2% of diet(see review by Apostolico, Vercillo, La Porta, &Ragni, 2016). Since ungulates are usually too large for awildcat to kill, it is likely that a large part of this food is
obtained through scavenging, but predation cannot be
excluded, because such dietary studies rely on analysis of
stomach content or scats and are thus not able to distinguish between predation and scavenging. Thus, the scavenging behavior and the ecological role of European
wildcats within the scavenging guilds in Europe remains
poorly understood. To fill this knowledge gap, we usedmotion-triggered camera traps deployed at 48 experimentally-set deer carcasses in forest ecosystems in Slovenia
and Italy to record scavenging by European wildcats. Our
goal was to describe the frequency of scavenging and557

caching behavior, observe potential interspecific interactions at the carcasses, and to study circadian and seasonaluse of the carcasses by the wildcats.

2 | METHODS
Our study was conducted in the forest landscape ofSouthern Slovenia and Northeastern Italy, where three
regions were selected to monitor scavenging activity on
experimentally-set ungulate carcasses: Kočevska(45 300 –45 490 N, 14 420 –14 600 E), Notranjska (45 350 –45 580 N, 14 150 –14 300 E) and Primorska-Kras (45 320 –46 040 N, 14 130 –14 500 E). Kočevska and Notranjska arepart of the Dinaric forest ecosystem characterized byhigh cover of mature forests dominated by fir and beech
associations. The average annual temperature in theseregions ranges from 6 to 9 C and snow cover lasts from50 days at 500 m and up to several months at higher elevations. Primorska-Kras is located near the SlovenianItalian border and has stronger Mediterranean influenceand lower forest cover with predominant oak, pine andhophornbeam associations. The average annual temperature in this region ranges from 9 to 13 C and snow
cover is mostly absent or lasts less than 10 days per year.
No reliable estimates of European wildcat densities exist
for these regions, but according to personal experiencesfrom the fieldwork in these regions, we assume thatdensities are higher in Dinaric forests than in the
Primorska-Kras region.
We placed camera traps on deer carcasses (n = 48)
killed through vehicle collisions in the three study areas:
24 carcasses of European roe deer (Capreolus capreolus)
and one carcass of red deer (Cervus elaphus) in Kočevska,
six roe deer and two red deer carcasses in Notranjska,
and 15 roe deer carcasses in Primorska-Kras (Table 1).
Carcasses were placed at randomly-selected locations
within the study area and monitored with several typesof motion-activated camera traps, including commercial
camera traps with black infrared (UOVision UV565GD,
Scoutguard Sg580m and U-Way U150X) in Kočevsko andPrimorska-Kras, and infrared-sensitive IP camera systemset (Mobotix M22M-Sec-Night) with independent infrared light (n = 7) or DSLR camera (Canon 350D) withwhite flash (n = 1) in Notranjska. Monitoring of carcasses
took place in 2010–2020 through all the seasons. All carcasses were monitored until all muscle tissue was consumed or/and completely decomposed.
We manually checked all the recordings andextracted those with wildcats. For the purpose of analysis,
we separated the year into two seasons: winter(December–March) and summer (April–November). We
calculated the proportion of carcasses used by the558

KROFEL ET AL.

SeasonKočevska

Notranjska

Primorska-Kras

All areas

Winter50% (4)25% (4)– (0)38% (8)

Summer0% (21)

0% (4)

0% (15)0% (40)All year

8% (25)13% (8)0% (15)

6% (48)T A B L E 1 Proportion of carcasses
scavenged by the European wildcats ineach study area and season

Note: Sample sizes are provided in the parenthesis. Seasonal differences were statistically significant(Fisher's Exact Test, df = 1, p = .008).

wildcats in each study area and each season, recorded
number of visits to the carcasses and time required forthe wildcats to find the carcasses, analyzed circadian
activity and noted feeding and carcass-caching behavior,as well as any interspecific interactions taking placearound the monitored carcasses. We did not attempt tomeasure the duration of visits, because in most cases theexact time of departure from the carcass was notrecorded. We counted recordings as different visits, when
they were more than 30 min apart without any record inbetween. Due to difficulties in distinguishing different
wildcats from the video recordings, the number of individuals using the carcasses remains unknown. In
Notranjska, the only carcass visited by the wildcat wasmonitored with a DSLR photo-camera with white flash,which resulted in a single photo. Possibly the flash disturbed the wildcat and because no video recordings wereavailable, we were not able to observe behavior in any
details. Therefore we used this record only for calculating
the proportion of carcasses found by the wildcats andtime needed for wildcats to find the carcasses. We usedkernel density estimation to determine the activity patterns of wildcat visits to carcasses, by transforming thetime each visit started into radians values. We then used
the overlap package (Meredith & Ridout, 2017) to fit thedata to a circular kernel density and estimated the activity among time periods from the distribution of the kernel density. We used Fisher's Exact Test to tests for
significance of seasonal differences in the use ofcarcasses.

3 | R E SUL T S
European wildcats were recorded on two (8%) of themonitored carcasses in Kočevska (both were from roedeer), one (13%) of the monitored carcasses in Notranjska(red deer), and none of the carcasses in Primorska-Kras(Table 1). All three instances of wildcat scavenging
occurred in winter, and scavenging was significantly
more often in winter than in summer (Fisher's Exact
Test, df = 1, p = .008). Carcasses were discovered by the
wildcats on average 13.4 days after deployment (on 2nd,
8th and 30th day after deployment). In total, we recorded

106 visits to the carcasses. In Notranjska, the wildcat visited the carcass only once (likely due to disturbance bycamera flash), while in Kočevska, wildcats visited carcasses on average 52.5 times per carcass (46 and 59 times
for each of the carcasses) with an average return interval
between consecutive visits of 6.2 h (n = 103) and maximum of 41 h (average for each of the carcasses: 4.4 and7.8 h). On average, the wildcats visited carcass 3.2-times
per day (average for each of the carcasses: 2.7 and4.2-times). The first carcass in Kočevska was recorded visited for 22 days (visited at least once on every day except
the ninth day) and the second carcass was visited for
11 days (visited every day).
We recorded wildcats feeding on these carcasses on
36% of the visits (n = 105; 35% for the first carcass and
37% for the first carcass). They were feeding predominantly on the hindquarters and afterwards to a lesser
extent on the abdomen, back, chest cavity, shoulders andneck (Supporting Information 1). Caching behavior wasobserved at both carcasses monitored with video surveillance, when wildcats used leaves, twigs or snow to partly
or completely cover the carcass with their front paws
(Supporting Information 2). Wildcats were recorded covering the carcass at least on 7% of the visits (n = 105; 5%
and 9% for each carcass) and covering was more frequent
during the first visits to each of the carcasses (57% of therecorded caching behaviors occurred during the first
2 days after wildcats discovered the carcasses). When
returning to the carcasses, wildcats sometimes used front
paws to again expose the carcass for feeding. The same
behavior was used when snowfall covered the carcasses.
In deep snow, wildcats also used carcass for resting by sitting or lying on top of them (Supporting Information 3).
We did not observe any urine-spraying or defecating by
the wildcats around the carcasses, but claw-marking(another scent-marking behavior) was recorded once
(Supporting Information 4). Wildcat visits to the carcasses were recorded throughout the 24-h cycle with acrepuscular pattern and the strongest peak in the earlymorning (Figure 1). Circadian use was similar for bothcarcasses (Supporting Information 5, Figure S1).
We recorded a confrontation with an European badger (Meles meles), which was aggressively chased fromthe carcass by the wildcat with raised tail, growling,

KROFEL ET AL.

F I G U R E 1 Circadian use of roe deer carcasses by theEuropean wildcats in Kočevska, Dinaric Mountains, Slovenia as
estimated via kernel density

hissing and snarling (Supporting Information 6). Twoother potential interactions were noted, but only the
wildcat could be seen on the video, while the other animals were outside of the camera view and could only beheard. Other scavengers recorded at the same carcasses
as wildcats, but which we did not observe to interact
directly with the wildcats, include (with % of all monitored carcasses that were visited by given species): red fox(Vulpes vulpes; 79%), wild boar (Sus scrofa; 29%), mouse(Apodemus sp.; 2%), common buzzard (Buteo buteo; 27%),
Ural owl (Strix uralensis; 2%) and common raven (Corvus
corax; 10%). Scavengers at other carcasses during thestudy included: brown bear (Ursus arctos; 42%), golden
jackal (Canis aureus; 2%), domestic dog (Canis familiaris;2%), beech marten (Martes foina; 38%) and great tit(Parus major; 2%).

4 | DISCUSSION
We have documented that European wildcats scavenged
from 38% of the carcasses we placed during winter, andthat they fed on these carcasses for extended periods withseveral visits per day. The extensive feeding, frequentcaching behavior and active defense from other scavengers indicates that the wildcats recognized the carcasses
as an important food source. Although the sample size
was relatively small (48 carcasses in total and 8 from winter period), this indicates that scavenging could be an
important, but neglected aspect of behavioral ecology ofEuropean wildcat during the winter period.
Many facultative scavengers use carrion as a supplemental food source (Wilson & Wolkovich, 2011). This isespecially true during winter when other food sources
may be less available but energetic needs are often higher(Krofel, 2011; Selva et al., 2005; Wilmers et al., 2003). For
European wildcats, large ungulate carcasses may be an
important source of food during winter because rodent

559

availability could be low because of snow cover (Selva
et al., 2005; Sonerud, 1986). Winter is also the period
when availability of ungulate carcasses is highest due toincreased natural mortality connected with high snow
levels, cold temperatures and reduced food availability
for herbivores (Selva et al., 2005; Selva, Jedrzejewska,
Jedrzejewski, & Wajrak, 2003). More frequent consumption of ungulates during winter by the European wildcats
was also suggested by the diet analyses (Apostolico
et al., 2016). Absence of wildcats at the monitored carcasses in Primorska-Kras region could be connected toseveral factors, but is most likely related to the lack ofmonitored carcasses during winter period in this region.
Many carnivores cache animal carcasses they havekilled or scavenged from as a means of food preservation.
Caching can be used for long-term storage of surplus foodduring times of hyperabundance, such as when arctic
foxes (Alopex lagopus) store waterfowl eggs from nestingseason for later consumption (Careau, Giroux, &Berteaux, 2007). But more often, caching is used forshort-term food storage, especially for large animal carcasses. For example, bears will cover the carcasses ofungulates or other large prey they kill or scavenge inorder to consume over a period of days (Allen, Wittmer,
Ingaki, Yamazaki, & Koike, 2020) and similar behavior isused by several felid species, especially for remains of theprey they have killed (Hunter, 2015). There is very limited literature available about the caching behavior ofEuropean wildcats, for example, Macdonald (1976)
reported probable caching in captive conditions andRuiz-Villar, López-Bao, and Palomares (2020) recently
reported about a wildcat scavenging and caching a roadkilled deer. Most of the previously documented incidents
actually refer to the African wildcats (L. Hunter, pers.
comm.), which were formerly treated as a subspecies ofF. silvestris, but are now recognized as a distinct species(Kitchener et al., 2017). However, we have shown that
European wildcats regularly cached both carcasses they
repeatedly scavenged from. They used leaves, twigs andsnow to cover the carcass (sometimes completely) in asimilar way as observed in felids that frequently consume
ungulates, for example, Eurasian lynx (Krofel et al., 2019)and puma (Bischoff-Mattson & Mattson, 2009). Thisbehavior is thought to effectively slow down decomposition and reduce probability of carcass detection by competing invertebrate and vertebrate scavengers (BischoffMattson & Mattson, 2009; Krofel et al., 2019). The
observed caching activity, along with the high frequency
and prolonged feeding, suggests wildcats are trying topreserve the carcass for as long as possible. We suggestthat caching behavior could be regularly performed by
European wildcats when using larger carcasses, but waspreviously likely neglected due to limited research effort

560

to document this behavior. Also during our monitoring,
we likely missed some of this behavior, because themoment of departure was often not recorded on the camera and wildcats mostly covered the carcasses before
leaving.
New knowledge about the ungulate-caching behavior by European wildcats has implications also forresearchers and management activities that use animal
signs to detect presence of wildlife species. Field guides
for animal tracking in Europe often suggest that caching of ungulate carcasses by covering with leaves, snow
and other material is a reliable sign of the lynx or bearpresence (e.g., Bang, Dahlstrøm, & Walters, 2001;
Krofel & Potočnik, 2016; Molinari, Breitenmoser, &Molinari, 2000). Our observations demonstrate thatpartly or completely covered ungulate carcass could
also be the result of scavenging by a European wildcat.
While caching by bears can be typically distinguishedby large amount of material used to cover the carcass,caching by the wildcat results in very similar signs tothose made by the Eurasian lynx. Besides, the observed
feeding pattern of the wildcats (predominant feedingon the hindquarters and gradual proceeding towards
the neck) was the same as reported in Eurasian lynx
(Krofel et al., 2019). Thus we suggest caution when
determining lynx kill sites only based on the signs ofcovering the carcass, in regions where wildcats arepresent.
Use of camera-traps also enabled us to observe interspecific interaction between European wildcat andEuropean badger. This was likely connected with theinterference competition for the carcass, as badgers wererecorded scavenging on this and other carcasses.
Although European badger is larger in body size compared to European wildcat, the latter managed to defend
the carcass during the encounter we observed. This could
reflect higher resource holding potential of the wildcat
compared to the badger, similar to other felid species thatare known for their higher resource holding potential
compared to other carnivores (Allen & Krofel, 2017).
Resource holding potential is often not dependent only
on body size, but can also be related to other physical features, behavioral traits, motivation and ownership status(Allen & Krofel, 2017).
The advent of camera traps has been already instrumental in documenting cryptic scavenging behaviors ofwild felids and other wildlife (e.g., Allen et al., 2015;
Krofel & Jerina, 2016; Peers, Majchrzak, Konkolics,
Boonstra, & Boutin, 2018). Using an experimental design
and monitoring of ungulate carcasses with camera traps
have enabled us to study this neglected aspect ofEuropean wildcat ecology. In this way we were able todifferentiate scavenging from predation, which is often

KROFEL ET AL.

difficult with the use of traditional methods, such as scator stomach content analysis, used in previous research.
The use of other field methods would also make it difficult to document carcass caching by wildcats, which islikely why it has gone largely undetected until now.
Therefore we recommend further research using thisapproach to better understand behavior and ecologicalprocesses connected with scavenging by wildcats andother felids, including interspecific interactions.
A C KN O WL ED G EME N T S
We thank U. Fležar, M. Hunjadi, L. Kozlan, T. Oliveiraand I. Smole for their help with the fieldwork, L. Hunterfor help with the literature search and two anonymous
reviewers for providing several useful suggestions on how
to improve the manuscript. This work was funded by the
Slovenian Research Agency (grants no. P4-0059 andN1-0163).
ORCID
Miha Krofel https://orcid.org/0000-0002-2010-5219
Maximilian L. Allen https://orcid.org/0000-0001-8976889X
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How to cite this article: Krofel M, Južnič D,
Allen ML. Scavenging and carcass caching
behavior by European wildcat (Felis silvestris).
Ecological Research. 2021;36:556–561. https://doi.
org/10.1111/1440-1703.12211