Wolf diet in an agricultural landscape of north-eastern Turkey

Abstract

We investigated wolf feeding ecology in Kars province, north-eastern Turkey, by analyzing 72 scat samples collected in spring 2013. Ongoing camera trap surveys suggest that large wild ungulates are exceptionally rare in the region. On the contrary, livestock is abundant. Accordingly, scats analysis revealed that livestock constituted most of the biomass intake for wolves, although small mammals were the most frequent prey items. Wild ungulates were occasional prey, and although wolves make use of the main village garbage dump as a food source, garbage remains were scarce in scat samples. Wolf dependence on anthropogenic resources, primarily livestock, generates human-wildlife conflicts in the study area. Uncontrolled carcass disposal seems to boost this wolf behaviour. Synanthropy enhances the probability of wolf-human encounters and thus increases the risk of direct persecution, vehicle collisions, and hybridisation with dogs. When livestock is not available, small mammals are an important alternative prey for wolves. This may increase interspecific competition, particularly with lynx, which is also lacking natural prey in the area. Our preliminary results contribute to wolf ecology and conservation in the Anatolian-Caucasian range, where further studies are urgently needed to generate baseline data.

Full text

Mammalia 2015; aop
*Corresponding author: Claudia Capitani, Environment Department,
University of York, Heslington, York, YO10 5DD, UK,
e-mail: claudia.capitani@york.ac.uk.
http://orcid.org/0000-0002-1899-8679
Mark Chynoweth: Department of Biology, University of Utah, 257
South 1400 East, Salt Lake City, 84112 UT, USA
Josip Kusak: Department of Biology, University of Zagreb,
Heinzelova 55, 10000 Zagreb, Croatia
Emrah Çoban: KuzeyDoğa Society, Ortakapi Mah. Sehit Yusuf Cad.
No:93 Kat:1, Merkez, Kars 36100, Turkey
Çağan H. Şekercioğlu: Department of Biology, University of Utah,
257 South 1400 East, Salt Lake City, 84112 UT, USA; KuzeyDoğa
Society, Ortakapi Mah. Sehit Yusuf Cad. No:93 Kat:1, Merkez,
Kars 36100, Turkey; and College of Sciences, Koç University,
Rumelifeneri, Sariyer 34450, Istanbul, Turkey
Short Note
Claudia Capitani*, Mark Chynoweth, Josip Kusak, Emrah Çoban and Çağan H. Şekercioğlu
Wolf diet in an agricultural landscape
of north-eastern Turkey
Abstract: In this study, we investigated wolf feeding ecol-
ogy in Kars province, north-eastern Turkey, by analysing
72 scat samples collected in spring 2013. Ongoing camera
trap surveys suggest that large wild ungulates are excep-
tionally rare in the region. On the contrary, livestock is
abundant. Accordingly, scats analysis revealed that live-
stock constituted most of the biomass intake for wolves,
although small mammals were the most frequent prey
items. Wild ungulates were occasional prey, and although
wolves make use of the main village garbage dump as a
food source, garbage remains were scarce in scat samples.
Wolf dependence on anthropogenic resources, primarily
livestock, generates human-wildlife conflicts in the study
area. Uncontrolled carcass disposal seems to boost this
wolf behaviour. Synanthropy enhances the probability
of wolf-human encounters and thus increases the risk of
direct persecution, vehicle collisions, and hybridisation
with dogs. When livestock is not available, small mam-
mals are an important alternative prey for wolves. This
may increase interspecific competition, particularly with
lynx, which is also lacking natural prey in the area. Our
preliminary results contribute to wolf ecology and conser-
vation in the Anatolian-Caucasian range, where further
studies are urgently needed to generate baseline data.
Keywords: generalist carnivore; human-wildlife conflict;
livestock scavenging; scats analysis.
DOI 10.1515/mammalia-2014-0151
Received October 14, 2014; accepted April 22, 2015
Wolves (Canis lupus Linnaeus, 1758) have been studied
throughout their distribution, but very little is known
about the Anatolian-Caucasian populations. Given its
ecological and geographical continuity with vast areas of
Central Asia and the Middle East, Turkey plays a central
role in maintaining wolf populations throughout the
region. The mountains of Turkey have served as a reser-
voir for the wolves surviving in Syria (Boitani 2003); yet,
habitat loss, illegal killing, taking pups from the wild and
vehicle collisions have resulted in a decline of wolves and
other large-carnivore populations in Turkey (Şekercioğlu
et al. 2011). Wolves are a species under protection from
hunting according to the Article 4 of Turkey’s Terrestrial
Hunting Law (Tuğ 2005), and the Ministry of Forestry
and Water Affairs is in charge for the management of this
species (Anonymous 2012). Monitoring the status of the
wolf population in Turkey is essential for the conservation
of the species both in the country and over a broader area.
The local environmental organisation KuzeyDoğa
Society (www.kuzeydoga.org), in collaboration with the
General Directorate of Nature Conservation and National
Parks, supported the creation of the first wildlife cor-
ridor in Turkey, eventually designated in 2011 with the
Ministry of Forestry and Water Affairs (Şekercioğlu 2012).
The corridor aims to connect isolated forest remnants
through reforestation, to provide habitat connectivity and
to facilitate the movements of large carnivores and their
prey species. It will cover 22,346 ha and will extend for
136 km, from Kars province, north-eastern Turkey, to the
extensive Caucasus forests on the Turkey-Georgia border.
For the corridor to be effective, it is critical to improve the
understanding of large-carnivore population dynamics
and spatial ecology in the area.
Opportunistic surveys over the last decade suggested
that primary prey species for wolf in Turkey are red deer
(Cervus elaphus Linnaeus, 1758), roe deer (Capreolus
capreolus Linnaeus, 1758), wild boar (Sus scrofa Linnaeus,
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2C. Capitani etal.: Wolf diet in north-eastern Turkey
1758), brown hare (Lepus europaeus Pallas, 1778) and live-
stock (Can O.E. personal communication, Anonymous
2012). However, quantitative investigations on wolf diet
and in particular on the relative share of wild and domes-
tic ungulates have not been conducted to date.
In this study, we present the results of a quantita-
tive assessment of wolf diet based on scat analysis con-
ducted around Sarikamiş, Kars (Figure 1). Our goals are
to contribute baseline data on large-carnivore ecology in
the extended wildlife corridor area and to improve the
general knowledge on wolf ecology in Turkey. The study
area (approximately 550 km2) is located on a high plateau
at the intersection of Caucasus and Irano-Anatolian global
biodiversity hotspots. Altitude ranges between 1900 and
3120 m asl. The landscape is characterised by patches
of forest spaced out by grassland. Although fragmented,
forests cover approximately 60% of the study area. Only
15% (49.7 km2) of the forested areas is included in the
Sarikamiş-Allahuekber Mountains National Park (hereaf-
ter SAM NP) (Figure 1).
Figure 1:Location of the study area: SAM NP and surrounding forest
in north-eastern Turkey. The SAM NP is fragmented forest in a land-
scape dominated by human activity, mainly livestock grazing. We
collected 72 wolf scats during a 1.5-month period in May–June 2013.
Forests consist almost exclusively of Scots pine (Pinus
sylvestris Linnaeus, 1753), while understory vegetation
is scarce, with consequent scarcity of food resources for
browsers.
Based on extensive camera trap surveys, wild boar is
present at low density, and roe deer is rare (Chynoweth
etal. unpublished data). On the contrary, livestock is abun-
dant. About 851,445 livestock heads have been registered
in the Kars province in 2012 (Ministry of Food, Agriculture
and Livestock, Republic of Turkey). Cattle (Bos taurus Lin-
naeus, 1758), sheep (Ovis aries Linnaeus, 1758) and goats
(Capra hircus Linnaeus, 1758) roam freely on pastures from
April to November in average climate conditions. Wolf, bear
(Ursus arctos Linnaeus, 1758) and lynx (Lynx lynx Linnaeus,
1758) are present in the area. At the time of this study, at
least two wolf packs occupied the area, and reproduction
was observed in one of them (Chynoweth etal. unpublished
data). The scarcity of natural prey species leads wolves, as
well as brown bears, to feed at garbage dumps and on live-
stock, increasing the human-carnivore conflict.
During 3 weeks between May and June 2013, we inten-
sively searched for signs of wolf presence and collected
scats over a 307 km network of forest roads (Figure 1).
We identified wolf scats on the basis of their size, shape,
content and smell (Jedrzejewski and Sidorovich 2010).
Scats of uncertain origin were discarded. Despite draw-
backs pointed out for scat analysis and the related prey
use indices (Klare etal. 2011), this methodology is helpful
in preliminary surveys of carnivores’ diet and is still
widely used, which facilitates comparisons with results
from different studies.
Given the limited sample size, we tested for adequacy
of sample effort by calculating the Brillouin diversity index
(Hass 2009), according to the equation ∑
=i
lnN!-lnn
Hb
N
where Hb is the diversity of prey in the sample, N is the
total number of individual prey categories in all samples
and ni is the number of individual prey in the ith category
(Brillouin 1956). An Hb diversity curve was calculated by
bootstrapping the sample 10,000 times with replacement
to obtain a mean Hb and 95% confidence interval, varying
the sample size from 2 to 100, in increments of 2. The Hb
increment curve was then calculated from the incremen-
tal change in each mean Hb with the addition of two more
samples. Adequacy of sample size was determined by
whether asymptotes were reached in both curves when
plotted against the sample size.
For every sample, the macroscopic components
(hairs, bones, hooves, claws, garbage remains, etc.) were
separated from the remaining matrix, and the volume of
each item was visually estimated to the nearest 5%. In
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C. Capitani etal.: Wolf diet in north-eastern Turkey3
other studies (Rigg and Gorman 2004, Tumanov 1998, Van
Duyne etal. 2009, Mattioli etal. 2011; Table 1).
Finally, we investigated the relationship between
diet composition and livestock availability on pastures.
In 2013, livestock was reported to begin grazing on pas-
tures from the third week of April. We estimated the depo-
sition time of our samples by collection date and degree
of degradation). Samples with an estimated deposition
time up to the third week of April were assigned to the
season when livestock is not available to predators (here-
after season A). Remaining samples were assigned to the
season when livestock are grazing on pastures and are
vulnerable to predation (hereafter season B). We com-
pared the use of two prey groups – specifically livestock
and small mammals – and grouped the main food catego-
ries to maintain adequate sample size and to account for
the undetermined categories within each group. We tested
for seasonal differences in the mean volume of each group
by applying the Wilcoxon rank-sum test.
Statistics were performed in R Core Team (2014). R:
A language and environment for statistical computing.
R Foundation for Statistical Computing, Vienna, Austria.
URL http://www.R-project.org/.
During the surveys, we collected 72 wolf scats useful
for diet analysis, whose deposition time we estimated to
vary from mid-March to mid-June. Fresh scats (deposi-
tion time < 2 days) were not collected for the diet analy-
sis since they were intended for other purposes. Both Hb
index mean and incremental change curves reached an
asymptote, and the incremental change declined below
1% at ≥ 36 samples (Figure 2), indicating that the sampling
effort was adequate.
Given information collected through camera trapping
(Chynoweth etal. unpublished data), we assumed that
most cases, mammal hairs were identified by examina-
tion of the medulla and cuticular surface structures under
a microscope and compared with a specific hair atlas
(Debrot et al. 1982). In few cases, hair and bones were
compared with reference collections and museum speci-
mens. For some samples, the species could not be deter-
mined, because of the poor quality of the remains or the
lack of specific reference material.
The utilisation rate of different food items was calcu-
lated by frequency of occurrence per item (hereafter FO)
and mean percent volume (hereafter MPV), following
previous studies (Ciucci etal. 1996, Capitani etal. 2004).
Remains contributing < 5% of the total scat volume were
considered as traces and not accounted for utilisation rate.
Utilisation indices, in particular FO, tend to underestimate
the share of big prey compared to the small ones and can
be misleading when prey greatly differ in size (Klare etal.
2011). Therefore, we applied a biomass model to convert
the equivalent number of scats in biomass and calculated
the relative share of prey categories. Biomass models are
sensitive to the weights of prey used during experimental
feeding trials (Klare etal. 2011); thus, we chose the model
which would cover the range of prey weights found in our
sample (Table 1). We used the equation developed by Floyd
etal. (1978), Y = 0.02X+0.038, where Y is the kilograms of
prey per collectable scat and X is the mean prey weight
(kg). We excluded garbage remains and undetermined
mammals that we could not estimate a mean weight. We
calculated the biomass using the prey weights reported in
Table 1:Composition of wolf diet in SAM NP and surrounding forest
in north-eastern Turkey, based on scats analysis (n=72; May–June
2013).
Food category MPV (%) FO (%) BM PW (kg)
Squirrel ....a
Cattle ...b
Hare ...
Small rodents ....
Sheep ...c
Undetermined livestock...c
Wild boar ....a
Undetermined mammal..––
Horse ...c
Bear ...d
Garbage ..––
Total 
Prey abundance is quantified by MPV, FO and biomass share (BM)
according to Floyd etal. 1978. aMean weight of wild boar was
estimated accounting for the weight classes identified in the scats,
following Mattioli etal. (2011). Mean weights of prey (PW) followed
previous studies: bRigg and Gorman (2004), cVan Duyne etal. (2009)
and dTumanov (1998).
Figure 2:The Brillouin diversity index (Hb) mean and 95% confi-
dence intervals (CI) and incremental change curves. Mean and 95%
CI were obtained by resampling with replacement 10,000 times.
Mean and incremental change curves reached an asymptote, and
the incremental change declined below 1% at ≥36 samples.
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4C. Capitani etal.: Wolf diet in north-eastern Turkey
prey availability varies little across the study area and
analysed the data cumulatively. Analysis of 72 scats docu-
mented a total of 80 food items, which were assigned to 11
food categories (Table 1). Only eight scats contained two
different food items at the same time, and so MPV and FO
resulted highly correlated (Spearman’ correlation index,
Rs = 0.98, p < 0.01).
The most frequent food categories were squirrel
(Sciurus vulgaris Linnaeus, 1758 and Sciurus anomalus
Gmelin, 1778) and cattle, followed by hare, sheep and small
rodents (Table 1), which included black rat (Rattus rattus
Linnaeus, 1758) and other undetermined rodent species.
Overall, small mammals were the most abundant in scats,
summing up to 45.2% of MPV. Livestock comprised 40.9%
of MPV, including horse and other undetermined livestock
remains, which most likely belong to either sheep or goat.
Wild boar remains were found in six samples only, including
one piglet. Though wolves have been frequently observed
feeding at the main dump site of the village (authors’ obser-
vations), garbage remains were rare; food remains taken at
the dump could be difficult to recognise unless associated
with undigested material. Finally, very exceptionally, one
scat contained hairs and a claw from a bear cub.
As expected, biomass shares largely differed from uti-
lisation indices (Table 1). Livestock represented 83.7% of
biomass, while small mammals share totally amounted to
10.8% (Table 1). The use of livestock and small mammals
differed between seasons A and B (Figure 3). Livestock
increased from 27.8% of MPV in season A to 54.1% of MPV
in season B (Wilcoxon test, W = 477.5, p = 0.031). On the con-
trary, small mammals decreased from 58.3% to 31.9% of
MPV (Wilcoxon test, W = 826.5, p = 0.025).
According to our results, Sarikamiş wolves have
a clear opportunistic feeding behaviour, using a wide
variety of food items but mostly relying on anthropogenic
resources, as found in other areas where wild prey are
scarce (see Meriggi and Lovari 1996, Peterson and Ciucci
2003 for a review). Furthermore, our data suggest a sea-
sonal variation in wolf diet due to the presence of free-
ranging livestock on pastures, a behaviour that has been
observed in other agricultural landscapes (Morehouse
and Boyce 2011). The scarce use of wild ungulates con-
firms that the density of these species is very low in the
study area. This scarcity of natural prey and the opportun-
istic feeding behaviour of wolves are probably leading to
trophic niche overlap with other carnivores, particularly
with lynx, which is also lacking natural prey in the area
(i.e., roe deer) and whose diet can be influenced by wolf
presence (Lelieveld 2013).
Our observations suggest that the wolves’ feeding
behaviour in Sarikamiş is related to local husbandry
Figure 3:MPV of prey found in 72 scats collected from SAM NP
and surrounding forest in north-eastern Turkey in season A (mid-
March to third week of April) and season B (fourth week of April to
mid-June). Livestock increased from 27.8% in season A to 54.1%
in season B (Wilcoxon test, W=477.5, p=0.031). Small mammals
decreased from 58.3% to 31.9% (Wilcoxon test, W=826.5, p=0.025).
practices. Open-air disposal of livestock carcasses to some
extent supports wolves that can scavenge on carcasses
when live prey is not available (Blanco and Cortés 2007).
These carcasses may also attract wolves to areas near live-
stock and could encourage livestock depredation (More-
house and Boyce 2011, Tourani etal. 2014). The authors
observed numerous openly disposed carcasses around
Sarikamiş area and once in broad daylight wolves could
be observed scavenging on a cattle carcass abandoned
on the roadside a few kilometres from Sarikamiş village.
Since conflicts are likely to be unevenly distributed across
the landscape, assessing local conditions of farms and
livestock husbandry practices is needed to provide spe-
cific mitigation tools (Rigg etal. 2011).
Synanthropy represents a major threat for wolves in
Sarikamiş because wolves are more likely to approach
human settlements to access trophic resources. This
enhances wolf-human encounters probability and results
in increased risks of direct persecution, vehicle collisions
(Fritts etal. 2003) and hybridisation with dogs (Kopaliani
et al. 2014). Human-induced mortality cases were often
reported in the study area (Chynoweth etal. unpublished
data), though detailed data on the wolf-livestock-human
dynamics are currently lacking.
As proposed for other areas where wolves largely
depend on anthropogenic resources, appropriate manage-
ment of garbage dumps and of livestock carcass disposal
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C. Capitani etal.: Wolf diet in north-eastern Turkey5
sites could reduce wolf-livestock conflicts and minimise
the chances of human-wildlife conflict and consequent
wolf mortality (Hosseini-Zavarei etal. 2013, Tourani etal.
2014). Such interventions should be realised in conjunc-
tion with actions for improving habitat suitability, for
example, the current efforts of the KuzeyDoğa Society to
improve habitat by increasing protected area coverage in
the region and to reforest the newly designated wildlife
corridor. Future efforts should also include management
of wild ungulate populations to increase the density of
wild prey, which could reduce livestock depredation to a
certain extent (Meriggi etal. 2011). As a potential solution
to mitigate human-wildlife conflict, wildlife managers
should consider reintroduction of native wild ungulates
(Meriggi and Lovari 1996), such as red deer that has been
reintroduced to other parts of Turkey (Gümüşhane Haberi
2013) and the Caucasus (World Wildlife Fund 2014).
Changes in socio-economic conditions could lead
to an alteration of wolf-prey dynamics in the study area,
where the number of livestock heads has dropped sharply
in the last decade (-78.5% goats, -15.6% cattle in Kars prov-
ince, source Republic of Turkey Ministry of Food, Agri-
culture and Livestock). In the overall region, cattle stock
dropped from about 15 million to 900,000 heads in Kars,
Igdir and Ardahan provinces in the past 50years (Nuri
Vatan, personal communication). Looking at future sce-
narios, continued abandonment of livestock husbandry
could exacerbate wolf-human conflict and, potentially,
cause a decline of the wolf population due to persecu-
tion and lack of prey. On the contrary, proper manage-
ment strategies could support an alternative scenario,
where abandonment of mountain areas by humans and
decreased grazing pressure by livestock would lead to the
increase of forest cover, wild ungulates and ultimately
biodiversity (Falcucci etal. 2007, Chapron etal. 2014).
The results of this study represent preliminary efforts
to investigate wolf ecology in the study area, though we
recognise that the low number of samples and the short
collection period could have biased our results. Further
investigations of year-round predator-prey dynamics,
local husbandry practices and interspecific interactions
are currently taking place. Long-term survey of wolf
ecology in the study area is required to design locally
tailored solutions to human-wildlife conflict, and, more
generally, it can contribute to the escalating debate on
large-carnivore conservation in human-dominated land-
scapes (Chapron etal. 2014).
Acknowledgments: We thank the General Directorate of
Nature Conservation and National Parks and Forestry
General Directorate of Turkey’s Ministry of Forestry and
Water Affairs for permitting our research, which was con-
ducted under the Eastern Turkey Wildlife Research and
Conservation protocol signed with the ministry. We thank
the Christensen Fund, National Geographic Society Edu-
cation Foundation, UNDP Small Grants Programme, the
University of Utah and the Whitley Fund for their support.
We are grateful to the KuzeyDoğa staff and volunteers for
their tireless efforts through the years and to the people
of Kars for their hospitality. We thank Dr. Luca Mattioli
for his contribution to prey remains identification and Dr.
Colin Courtney Mustaphy for commenting on a draft of
the original manuscript.
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