Animal Conservation.

Print ISSN 1367-9430

One way or another: predictors of wolf poaching in a

legally harvested wolf population
J. Suutarinen1 & I. Kojola2
1 University of Oulu, Oulu, Finland
2 Natural Resources Institute Finland, Rovaniemi, Finland

Keywords
Canis lupus; gray wolf; human–carnivore
conflict; human-caused mortality; poaching;
illegal killing; carnivores.
Correspondence
Johanna Suutarinen, University of Oulu,
Paavo Havaksen tie 3, 90014 Oulu, Finland.
Email: johanna.suutarinen@luke.fi
Editor: Julie Young
Associate Editor: Vincenzo Penteriani
Received 22 August 2017; accepted 13
February 2018
doi:10.1111/acv.12409
Abstract
Poaching is a major threat to large predator populations, but the predictors of poaching
are poorly mapped in developed countries where illegal killing is motivated by social
reasons. Poaching can be common although the species is legally hunted, such as in
the case of the wolf (Canis lupus). Our goal was to identify crucial motives of poach-
ing to find possible solutions to the ongoing wolf conflict. We studied predictors of
poaching on two spatial scales – countrywide (76 wolves) and territory [30 Global
Positioning System (GPS)-wolves] – during 2001–2016 in Finland. The countrywide
factors predicting illegal kill were as follows: (1) lifestage, with adult wolves showing
a remarkably high probability of being illegally killed in comparison with juveniles;
(2) the number of wolves killed legally in the local scale, that is, licensed wolf hunting
at the local scale decreased the likelihood of illegal killing, as did the total number of
legally hunted wolves; (3) total legal bag in the whole country; and (4) density of the
local human population, that is, low human density increased the probability of illegal
kill. For breeding adult GPS-collared wolves at the territory level, there was a positive
relationship between the tendency to cross roads and likelihood of being illegally
killed. Our results provided evidence that poaching is a matter of local intolerance
toward wolves and that the problem is mainly related to wolf hunting. Legal hunting
might decrease poaching, but seems inefficient as a long-term solution. To maintain a
viable wolf population, the poaching risk of breeding adults should be decreased. Pre-
dicted poaching probabilities could be used to tackle poaching in a preventive manner
by involving both decision makers and local residents in anti-poaching actions.

The negative attitudes and fear toward wolves (Treves,

Introduction
Poaching is a many-sided global issue, from large-scale illicit
trade to sporadic hunting offenses (Dickman, 2010; Challen-
der & MacMillan, 2014; Chapron & Treves, 2016; Pohja-
Mykr€a, 2016). In addition to habitat destruction in human-
altered landscapes (Ripple et al., 2014), poaching threatens
many wildlife populations (Liberg et al., 2012; Maisels
et al., 2013; Challender & MacMillan, 2014; Sharma et al.,
2014). Top predators are especially prone to conflicts due to
their wide home ranges and carnivorous behavior (Graham,
Beckerman & Thirgood, 2005; Treves & Bruskotter, 2014).
For example, in northern latitudes, poaching motivation
arises from carnivore-livestock and carnivore-game conflicts
(Graham et al., 2005; Andren et al., 2006; Persson, Ericsson
& Segerstr€om, 2009; Treves & Bruskotter, 2014) and is
intensified by human–human conflicts between the different
interest groups (Dickman, 2010; Pohja-Mykr€a, 2016).
With most controversial species, such as the gray wolf
(Canis lupus), the principal objective of illegal killing is to
reduce or remove the species (Treves & Bruskotter, 2014).
Naughton-Treves & Shelley, 2013; Dressel, Sandstr€ om &
Ericsson, 2015) appear to be due to a hyper-awareness of
the risks caused by these potentially risky animals (Dickman,
2010). Wolves cause livestock damage (Kaartinen, Luoto &
Kojola, 2009) and the loss of hunting dogs (Kojola et al.,
2004), as well as hunt moose (Alces alces), which is both
the primary prey for wolves and is considered to be the most
valuable game species in boreal regions (Timmermann &
Rodgers, 2005).
Illegal killing may threaten the recolonization of wolf pop-
ulations (Liberg et al., 2012; Olson et al., 2014; Chapron &
Treves, 2016; Suutarinen & Kojola, 2017). Poaching can
have a remarkable effect on both the population dynamics
and on individual behaviors of a social, monogamous group
living species (Borg et al., 2015), especially if breeding adult
wolves are exposed to high poaching risk (Suutarinen &
Kojola, 2017). Death of one or both breeding adults can
increase the risk of mortality of the offspring (Brainerd
et al., 2008), cause pack dissolution (Borg et al., 2015), and
decrease hunting success (Sand et al., 2006). Consequently,

414 Animal Conservation 21 (2018) 414–422 ª 2018 The Zoological Society of London
J. Suutarinen and I. Kojola
poaching has the potential to work in a super-additive way,
affecting the population to a greater extent than most other
types of mortalities (Milleret et al., 2016). Wolf poaching is
mostly unreported and difficult to prove, and therefore called
‘cryptic poaching’ (Liberg et al., 2012), resulting in an
unknown size and frequency of this type of crime (Biderman
& Reiss, 1967; Wellsmith, 2011). Due to cryptic poaching,
anti-poaching actions and achievements by authorities often
remain abstract.
Finland was recolonized by wolves in the 1990s (Kojola
et al., 2014). Since then, the population has been fluctuating
(Jansson et al., 2012; Kojola et al., 2014). According to the
Natural Resources Institute Finland (Kojola, 2005; Kojola
et al., 2014), the wolf population increased from c. 130 to
160 wolves from 2000 to 2003 up to 180–250 wolves from
2004 to 2008, with minor changes in between; then, the
population decreased to 120–160 wolves in 2009–2013 and
increased to 220–270 in 2014–2015, with a recent decline to
150–180 wolves (Natural Resources Institute Finland,
2017a). Annual reports by the institute include comprehen-
sive description of pack size and territory area locations,
which are based on observations collected from the hunters,
snow-tracking, noninvasive genetics, and positioning data of
the collared wolves (Natural Resources Institute Finland,
2017a).
The wolf’s legislative status in Finland is a protected
game species according to national legislation (Hunting Act
and related decrees) and regulations by the European Union
(EU, Habitats Directive, Council Directive 92/43/EEC). Wolf
hunting is license-based, and annual hunting quotas are
issued by Ministry of Agriculture and Forestry. Wolf man-
agement policy has undergone several changes in 2000–
2016. In 2015, the ministry launched a territory-based ‘toler-
ance hunting’, a hunting policy permitting hunting licenses
for management purposes (Ministry of Agriculture and For-
estry, 2015). These licenses were targeted on local wolf
packs to decrease wolf conflict in the territory areas.
Lethal control is often used as a tool for large carnivore
management besides sports hunting to maintain populations
at stable levels, to reduce conflicts over property, and to
build public support (Treves, 2009). In Finland, legal hunting
has been the administrative solution to ease discomfort
toward wolves and an attempt to reduce illegal killing (Min-
istry of Agriculture and Forestry, 2015). Epstein (2017)
describes such ‘tolerance hunting’ as hunting premised on
the hypothesis that the negative attitudes toward wolves that
lead to illegal killing will be ameliorated to the point of tol-
erating a greater wolf presence if people are allowed to leg-
ally kill wolves.
Although lethal control is used to tackle poaching in Uni-
ted States and Northern Europe (Chapron & Treves, 2016;
Epstein, 2017), contradictory arguments have been presented,
that is, there is no scientific support for the hypothesis
(Epstein, 2017), that lethal control does not inevitably
increase feeling of stewardship over large carnivores (Treves,
2009) or tolerance (Browne-Nu~nez et al., 2015) and that it
may impress a negative message about the value of wolves
(Chapron & Treves, 2016; but see also Pepin, Kay & Davis,
Animal Conservation 21 (2018) 414–422 ª 2018 The Zoological Society of London
Predictors of wolf poaching
2017). In addition, hunting may reduce the ecological func-
tion of these apex predators (Ordiz, Bischof & Swenson,
2013).
Here, we examined the predictors of poaching in a wolf
population that lives in a human dominated landscape and
has been exposed to various levels of legal harvest. We
aimed to predict poaching probabilities for different scenarios
of legal harvest.
Materials and methods
The Finnish landscape is dominated by forests: three-quarters
of the land area is covered by forests and one-tenth by agri-
cultural land (Statistics Finland, 2014). The majority of the
human population lives in urban areas, and the average
human density in Finland is 18 inhabitants per km2 (Official
Statistics of Finland, 2017). Finland has c. 300 000 hunters
out of the total population of 5.5 million people (Natural
Resources Institute Finland, 2017b). Calculated based on the
above statistics, the proportion of hunters ranges from <2%
in the Uusimaa province where the capital city Helsinki situ-
ates, up to 20% in the Kainuu province in east-central Fin-
land. Human density in our study area was c. 1.6 humans
per km2 and forest road density c. 1.2 km km 2.
We equipped 146 wolves with very high frequency (Telo-
nics, Mesa, AZ, USA) or GPS (Televilt, Lindesberg, Swe-
den; and Vectronic, Berlin, Germany) collars during 1998–
2016 (Kojola et al., 2006). Of these, we selected wolves to
be included in the analysis using the following criteria: (1)
when the death was caused by illegal killing or legal hunting
(Suutarinen & Kojola, 2017); (2) death occurred outside of
the reindeer herding area, where the wolf hunting policy is
more liberal than in the rest of Finland in order to keep the
reindeer herding area free of resident wolves (Kojola et al.,
2009; Ministry of Agriculture and Forestry, 2015); (3) indi-
viduals were not known to be a members of the same pack
as any other study animal included in the analysis; and (4)
they were able to be identified as juveniles (<2 years old) or
adults (>2 years old) based on their social status (Kojola
et al., 2006) or age defined from a tooth sample (Matson’s
Laboratory LLC, Manhattan, MT, USA, see also Ballard,
Matson & Krausman, 1995). Following the above criteria,
we chose 38 illegally and 16 legally killed collared wolves
that died during the calendar years 2001–2016.
To obtain the same number (n = 38) of legally and ille-
gally killed wolves for the analysis, we randomly sampled
22 legally killed wolves from the inclusive bag record data
of wolves that were legally hunted in 2001–2016 that met
the same criteria (n = 129). We conducted sampling using
the sample function in the program R (R Core Team 2015).
Kill site coordinates for legally shot animals were reported
by hunters as routine protocol. Sites for illegal kills were
assessed by using: (1) the exact kill site coordinates given in
a report (n = 2); (2) last position of the wolf (n = 8); and
(3) central spot of the territory area (n = 18), which was
measured as a 100% Minimum Convex Polygon (MCP) of
the GPS position. The kill site was known by its name in 10
illegal cases. We searched the coordinates for these place
415
Predictors of wolf poaching
names using a free online mapping application ‘Map Site’
by National Land Survey of Finland (https://asiointi.maanmit
tauslaitos.fi/karttapaikka/?lang=en).
Using the kill site as the center point, we created a circu-
lar range of 1000 km2 to calculate the spatial variables. The
wolf territory size is c. 1000 km2 in Finland, but because of
the variability in the size of the 100% MCPs (621–
3868 km2, Kaartinen, Antikainen & Kojola, 2015; see also
Mattisson et al., 2013), we used these computational ranges
in the country-level model (Mattisson et al., 2013; Ordiz
et al., 2015; Milleret et al., 2016).
The country-level model included 13 independent vari-
ables (Table 1). Spatial data were treated using ArcMap
10.3.1 (ArcGIS; ESRI, Redlands, CA, USA) and the
Geospatial Modelling Environment (Hawthorne L. Beyer,
Spatial Ecology LLC).
To examine the predictors in more detail, we used data
derived from countrywide data for a territory-level inspec-
tion, including only GPS-collared breeding adult wolves
(n = 30, Table 2). The territory-level model included two
independent variables which were selected on the basis of
the country model results (Table 3). We defined the territory
area as the 100% MCP and calculated the spatial variables
using this area.
We analyzed the data with logistic regression using a gen-
eralized linear model (GLM) with a binomial-dependent vari-
able (0 = legal, 1 = illegal). We standardized the explanatory
variables by scaling the means to 0 with a variance of 1 to
improve the interpretability of the regression coefficients
(Schielzeth, 2010). We tested correlation between continuous
independent variables using Spearman’s rank correlation test.
Model selection was made based on corrected Akaike’s
information criterion (AICc; Burnham & Anderson, 2002).
We used R package ‘AICcmodavg’ for Akaike calculations
(Mazerolle, 2017).
We created three a priori alternative models (Table 2),
which were based on the working hypotheses: (1) poaching
is related to legal wolf harvest, (2) poaching is related to
hunting in broader sense because wolves are considered
harmful for hunting and game management, and (3) poach-
ing is related to overlapping home ranges of wolves and
humans. We simplified a priori full models based on the
standard errors and confidence intervals of the independent
variables until we reached the smallest AICc per a priori
model. Then, we combined the simplified models with small-
est AICc and created a full combined model. We simplified
the combined model until reaching the smallest AICc. This
procedure resulted in 15 candidate models (Table 4a) which
we ranked based on their relative differences (Di AIC) and
strengths of evidence (Akaike weights xi). For territory-level
model, we created three alternative models: one with both
variables and both separately, and compared their Akaike
values (Table 4b).
We made multimodel inference on the variables ‘road
density’ and ‘legal bag change’ using model averaging and
resulting conditional standard error and confidence intervals.
We tested the effect of the possible outliers by making a
subset of the data excluding the high leverage point(s) for
416
J. Suutarinen and I. Kojola
each independent variable and examined with single-variable
GLM whether or not the exclusion influenced the direction
of the effect.
Logistic regression coefficients can be converted into odds
ratios (OR) which can be used to predict the probability of an
event to occur for a given set of predictor values. The number
of illegally killed wolves (MP) can be derived from the pre-
dicted probabilities as MP = N 9 mP, where N is the popula-
tion size and mP is the probability to be illegally killed. We
calculated the predicted number of illegal kills for a hypotheti-
cal population of 250 wolves with the approximate proportion
of adults and juveniles as in Finland (I. Kojola, unpubl. data):
adults represented 32% and juveniles 68% of the population,
that is, 80 adults and 170 juveniles. We calculated the predicted
poaching probabilities for adults and juveniles separately based
on the top-ranked model and used the original unstandardized
values of the independent variables.
Results
The likelihood of illegal kill at the country level was
affected by ‘legal bag range’, ‘total legal bag’, ‘human den-
sity’ and lifestage (Table 4a). At the territory level, the like-
lihood of illegal kill was predicted by the frequency that the
wolves crossed roads (Table 4b).
According to the top-ranked model, lifestage had the
strongest effect on the likelihood of illegal kill. The risk of
being illegally killed instead of legal kill was c. 5% for juve-
niles in comparison to adult wolves (juvenile vs. adult:
b = 2.87, SE = 0.78, OR = 0.06).
For a harvest strategy with a quota of 30 wolves (‘total legal
bag’ = 30) kept constant over the years (‘legal bag
change’ = 1) and distribution of one permit per local area (‘le-
gal bag range’ = 1) in the absence of human residence in the
area (‘human density’ = 0), the probability of being illegally
killed was 0.46 for adults and 0.05 for juveniles. For a more
populated area (‘human density’ = 10), the probabilities were
0.14 for adults and 0.01 for juveniles. For the hypothetical pop-
ulation of 250 wolves, the example with zero human density
would double the number of harvested wolves: 36 illegally
killed adults and 8 illegally killed juveniles, but the example
with higher human density would halve it: 11 illegally killed
adults and 2 illegally killed juveniles.
‘Legal bag range’ had a negative effect on the likelihood
of illegal kill. The likelihood of being illegally killed was
three times less likely if there was another wolf shot with
permission within the range during the hunting season in
comparison with the absence of licensed wolf hunting
(b = 1.26, SE = 0.55, OR = 0.28). A larger total number of
legally killed wolves (‘total legal bag’) also decreased the
likelihood of illegal kill (scaled variable b = 1.43,
SE = 0.57, OR = 0.24). For the unstandardized values, the
chance of illegal kill decreased by an odds ratio of 0.90
[95% CI (0.81, 0.96)], for an increase of one wolf in the
total legal bag, that is, c. 10% higher chance for the wolf to
die in a legal hunt (1/OR = 1.11). In terms of probabilities,
the poaching probability for adult wolves for a hunting quota
of 20 wolves would be 0.71, but for 40 wolves, the
Animal Conservation 21 (2018) 414–422 ª 2018 The Zoological Society of London
J. Suutarinen and I. Kojola Predictors of wolf poaching

Table 1 Country-level a priori models and independent variables of the illegality of a wolf kill model for wolves in Finland

Predictor Determination Key figures (unconverted)

(1) A priori model ‘legal wolf hunting’
Legal bag range Number of legally hunted wolves within the artificial 1000 km2 range Range: 0–5
during the hunting year the individual died, excluding the individual 1st–3rd quartile: 0–1
itself Median: 0
SD: 1.1

Legal bag region Number of legally hunted wolves within a game administration region of Range: 0–17
the Finnish Wildlife Agency (corresponding to the provinces) during the 1st–3rd quartile: 1–8
hunting season the individual died, including the individual itself Median: 4.5
SD: 5.2

Total legal bag Number of legally hunted wolves in the whole country during the Range: 7–70
hunting year the individual died, including the individual itself 1st–3rd quartile: 17–34
Median: 21
SD: 13.2

Legal bag change Change in total wolf bag in two consecutive hunting years when t + 1 is Range: 0.28–8.3
the year the individual died 1st–3rd quartile: 0.81–2.18
Median: 1.65
SD: 1.8

(2) A priori model ‘hunting versus wolf presence’

Moose bag change Change in moose bag within the game administration region, in two Range: 0.51–1.45
consecutive years, when Ht + 1 is the hunting year (Ht) the wolf died. 1st–3rd quartile: 0.83–1.12
Moose bag records by Finnish Wildlife Agency Median: 0.98
SD: 0.23
Hunter proportion Proportion of hunters of the total human population within the province. Range: 0.02–0.22
Number of hunters determined as the total number of people who had 1st–3rd quartile: 0.11–0.19
paid for the game management fee for the year that the wolf died. Median: 0.12
Human population data by Statistics Finland SD: 0.06
Dog damages Number of dogs killed by wolves according to the records of Finnish Cases (illegals):
Wildlife Agency. Data pooled over the years 2010–2015 (N = 213). Low: 16 (4)
Number of damages per province categorized to low (<10), moderate Mod: 2 (2)
(10–20) and high (>20) damage levels High: 58 (32)
Road density Road density (km km 2) of unpaved secondary roads (i.e., forest roads) Range: 0.35–2.58
within the artificial 1000 km2 range. Road data by National Land Survey 1st–3rd quartile: 1.0–1.53
of Finland Median: 1.17
SD: 0.42

(3) A priori model ‘overlapping home ranges’

Human density
Livestock damages
Population increase
Human population density (permanent residence) per km2 within the Range: 0.04–74.8
artificial 1000 km2 range in the year the wolf died. Human population 1st–3rd quartile: 0.56–5.1
density data by Statistics Finland Median: 1.63
SD: 13.1
Number of livestock damage events caused by wolves according to the Cases (illegals):
records of Finnish Wildlife Agency. Data pooled over the years 2010– Low: 11 (6)
2015 (N = 130 events, total 441 animals). Number of damages per Mod: 33 (15)
province categorized to low (<5), moderate (5–10), and high (>10) High: 32 (17)
damage levels
Rate of population increase, when n + 1 is population size in the year the Range: 0.67–1.48
individual died. Population sizes are based on estimates of population 1st–3rd quartile: 0.83–1.14
size in the end of the each year (see Suutarinen & Kojola, 2017) Mean: 1.01
SD: 0.18

Lifestage Adult > 2 years of age See Table 2

Juvenile < 2 years of age
Sex Female, male

Animal Conservation 21 (2018) 414–422 ª 2018 The Zoological Society of London 417
Predictors of wolf poaching J. Suutarinen and I. Kojola

Table 2 Description of the wolf data used in this study (b = 1.77, SE = 0.81, OR = 5.89 [95% CI (1.5, 40.5)]. ‘Yard
Adults Juveniles Total
visits’ did not effect on the likelihood of illegal kill (in terri-
Countrywide (females) (females) (females) tory-level full model b = 1.13 [95% CI ( 0.21, 4.68)],
SE = 1.17). ‘Road crossings’ model had the smallest AICc
Legal 19 (9) 19 (12) 38 (21)
value and Akaike weight (Table 4b).
Illegal 33 (14) 5 (1) 38 (15)
Total 52 (23) 24 (13) 76 (36)
Predicted poaching probabilities could be used to quantita-
tively estimate poaching levels in the following way: with a
Territory level Females Males Total hunting quota of 50 wolves (‘total legal bag’ = 50) put into
Legal 3 5 8 practice by removing one wolf per pack or local area (‘legal
Illegal 8 14 22 bag range’ = 1), and with the average human density within
Total 11 19 30 the 1000 km2 ranges (1.63 inhabitants per km2), the pre-
dicted probability of illegal kill would be 0.2 for adults and
The countrywide data were a sample of 76 wolves killed outside
0.01 for juveniles. For the hypothetical population of 250
the reindeer herding area in Finland 2001–2016. The territory-level
wolves, that would mean 16 illegally killed adults and 2
data were a sample of 30 breeding adult Global Positioning
juveniles. The total loss including both the legal and illegal
System-collared wolves killed outside the reindeer herding area in
harvests would be 68 wolves.
Finland 2003–2016.

poaching probability would decrease to 0.22 (‘human den- Discussion

sity’ = 0, ‘legal bag range’ = 1, ‘legal bag change’ = 1).
‘Legal bag change’ increased the likelihood of illegal kill
(b = 1.62, SE = 0.61, OR = 5.0). ‘Legal bag change’ had four
influential outliers, all of which were illegally shot in the same
province during the hunting season of 2000–2001. That hunt-
ing season had remarkably larger total legal bag than the sea-
son before, resulting in a change of 8.33. GLM with ‘legal bag
change’ as the single one independent variable resulted in
b = 0.08, but excluding the influential outliers b = 0.09. To
avoid erroneous interpretation, we held the ‘legal bag change’
at value 1 in all predicted probabilities.
The top-ranked model was only 1.96 times better than the
second-ranked model that included additionally ‘road den-
sity’ variable. The weighted average of the ‘road density’
estimate was 0.43 [unconditional SE = 0.42, 95% CI
( 1.25, 0.4)]. Confidence interval of ‘road density’ included
zero indicating that road density did not affect the likelihood
of illegal kill at the country level.
Based on the evidence that, at the country level: (1) ‘hu-
man density’ affected the illegality and (2) the effect of
‘road density’ was slightly unclear, we examined the effect
of secondary roads and human settlement within the territory
to the likelihood of illegal kill in more detail at the territory
level. Frequent crossing of roads while moving within the
territory increased the likelihood of being illegally killed
Our results speak of intolerance toward wolves and empha-
size the local focus on the wolf conflict. The results sup-
ported the assumption that legal hunting decreases illegal
killing. The higher poaching rates during shortages of legal
hunting permits might be an act of frustration and lead to
the removal of wolves in a one way or another. Despite
annual legal harvesting, the human–wolf conflict in Finland
has not led to coexistence, and tolerance for wolves has
remained low (Bisi et al., 2010; Pohja-Mykr€a, 2016).
Breeding adults suffered a very high poaching likelihood
in our study area, which represents a landscape of high
human impacts on forest structure and wildlife populations,
but low human density. In Finland, poaching has focused on
breeding adults with highest poaching risk in January–Febru-
ary (Suutarinen & Kojola, 2017), that is, simultaneously with
the breeding season. Adult wolves are essential for the conti-
nuity of a wolf population. Successful breeding of local wolf
packs in consecutive years could work as an indicator of an
absence of poaching. Securing the survival of breeding
adults could also increase local acceptance in the long run,
as wolves learn to avoid human settlements as they become
familiar with their territory (Kojola et al., 2016). In addition,
pack dissolution reduces a pack’s hunting success (Sand
et al., 2006) and can increase damage to domestic animals
(Wielgus & Peebles, 2014; Fernandez-Gil et al., 2016).

Table 3 Independent variables at the territory level of the illegality of a wolf kill model for wolves in Finland

Predictor Determination Key figures (unconverted)

Yard visits Proportion of GPS positions closer than 150 m from a house or other Range: 0.0005–0.9
building (Npos = <150 m) out of all GPS positions (Npos). 1st–3rd quartile: 0.004–0.1
Building location data by National Land Survey of Finland. Median: 0.006
SD: 0.02
Road crossings Proportion of road crossings measured as intersects of lines of Range: 0.2–1
consecutive GPS positions with roads inside the territory area 1st–3rd quartile: 0.3–0.5
(100% MCP). Road data by National Land Survey of Finland. Median: 0.41
SD: 0.17

GPS, Global Positioning System; MCP, Minimum Convex Polygon.

418 Animal Conservation 21 (2018) 414–422 ª 2018 The Zoological Society of London
J. Suutarinen and I. Kojola Predictors of wolf poaching

Table 4 Model ranking of the (a) countrywide and (b) territory-level illegality of a wolf kill models based on corrected Akaike’s information
criterion (AICc) of the logistic regression models with binary response variable (0 legal, 1 illegal)

K AICc Di AICc xi LL
(a) Countrywide models
Legal bag range + total legal bag + legal bag change + human density + lifestage 6 74.53 0.00 0.66 30.65
Legal bag range + total legal bag + legal bag change + road density + human density + lifestage 7 75.87 1.35 0.34 30.11
Legal bag range + total legal bag + human density + lifestage 5 84.58 10.05 0.00 36.86
Human density + lifestage 3 92.33 17.80 0.00 43.00
Human density + lifestage + sex 4 94.14 19.62 0.00 42.79
Lifestage 2 97.00 22.47 0.00 46.42
Human density + livestock damages + lifestage + sex 6 98.56 24.03 0.00 42.67
Legal bag range + total legal bag + legal bag change 4 98.83 24.30 0.00 45.13
Road density 2 99.07 24.54 0.00 47.45
Hunter proportion + road density 3 100.74 26.21 0.00 47.20
Human density + livestock damages + population increase + lifestage + sex 7 100.82 26.30 0.00 42.59
Legal bag range + legal bag region + total legal bag + legal bag change 5 101.00 26.47 0.00 45.07
Moose bag change + hunter proportion + road density 4 102.93 28.40 0.00 47.18
Moose bag change + dog damages + hunter proportion + road density 6 105.37 30.84 0.00 46.08
Legal bag range + total legal bag 3 106.16 31.64 0.00 49.92
(b) Territory-level models
Road crossings 2 31.94 0.00 0.50 13.75
Yard visits + road crossings 3 32.03 0.09 0.48 12.55
Yard visits 2 37.77 5.83 0.03 16.66

K stands for number of parameters in the model; Di AICc for AICci AICctop-ranked; xi for Akaike weight; and LL for log likelihood. Descrip-
tions of a priori models and independent variables are given in Tables 1 and 2.

Damages to domestic animals did not affect the poaching
likelihood, even though damages increase the negative atti-
tude toward wolves (Treves, 2009; Bisi et al., 2010). Dam-
ages to livelihood and property are often the basis for a
legal wolf hunting permit, and therefore, the effect on the
illegality can remain absent even though damages undoubt-
edly increase the mortality risk itself. Territory-level inspec-
tion of the effect of damages would demand detailed
information on the damage and the wolf kill, and might not
be even justifiable: such information would require large
dataset to fade out any identifiable information of personal
details, especially in the sensitive context of illegality.
Wildlife crime is victimless, usually with no witnesses,
and often takes place in remote areas. Risk of getting caught
by law enforcement is low in such offenses, which was sup-
ported by our results of higher human density decreasing
poaching likelihood. This highlights the poaching hotspots in
more remote areas, but can also be influenced by the possi-
ble bias caused by better chances of getting legal permits to
hunt wolves in more human populated areas. Nevertheless,
predicted poaching probabilities could be implemented by
law enforcement in allocating resources and in targeting
surveillance on areas with higher poaching risk.
An increased total legal bag increased the likelihood of
poaching, which indicates that swift changes in wolf hunting
might result in higher poaching rates. The effect was unclear
due to outliers, but should not be totally neglected. An
increased poaching rate due to an increased total legal bag
could also be a type of hyper-aware response to the indirect
message taken from the increased total legal bag – that the
wolf population requires greater harvest. To track the
consequences of management changes in depth, wolf man-
agement requires multidisciplinary inspection (Treves et al.,
2017) and straightforward interpretations that granting legal
hunting of wolves reduces poaching should be done with
caution.
The only significant predictor of illegal kill at the territory
level was the frequency at which wolves crossed roads.
Roads have been observed to increase human-caused mortal-
ity of wolves (Person & Russell, 2007). Finland has a dense
forest road network, which makes territories easily accessible
year round. The local wolf population is often under frequent
tracking from the roads by local hunters: for example, obser-
vations of wolf tracks are also collected nationally in Finland
as a part of wolf monitoring; for instance, the estimate of
the wolf population size in March 2017, of c. 150–180
wolves, was partially based on more than 3300 observations
from local hunters.
The countrywide and territory-level models offer supple-
mentary insight. The rate of population increase had no
effect on poaching likelihood, but road crossings indicating
the detectability of the local wolf population increased this
likelihood. Yard visits did not affect the poaching likelihood,
but higher density of human population decreased it. Local
and countrywide hunting measures were significant, but
regional measures were not. Therefore, it seems that from
the viewpoint of the local people, the subjective impression
of the number of wolves is more important than the actual
population size, further underlining the importance of attitude
and tolerance with regard to coexistence with wolves.
Higher level of legal hunting decreases the availability of
wolves for poachers. Increased hunting total legal bag might

Animal Conservation 21 (2018) 414–422 ª 2018 The Zoological Society of London 419
Predictors of wolf poaching
decrease poaching by simply decreasing the available catch,
and instead of solving the problem would end up just clean-
ing up the numbers.
Illegal killing of wolves is characteristically cryptic (Lib-
erg et al., 2012), and reported illegal kills are only the tip of
the iceberg. Therefore, correct in-time measurements of such
hidden criminality are difficult, if not impossible, to obtain.
Numerical estimates are required to reliably measure whether
higher levels of legal hunting diminishes illegal killing of
wolves or not, which factors predict poaching, and at which
scale the policy actions are most efficient, especially consid-
ering the controversial ‘tolerance hunting’ (Treves, 2009;
Browne-Nu~nez et al., 2015; Chapron & Treves, 2016;
Epstein, 2017). Legal hunting can act as a short-time buffer
against illegal killing. A zero-hunting policy could be detri-
mental for wolves and their conservation because of the
ongoing conflict. Zero-hunting policy would probably not
gain public support especially in a country such as Finland
where hunters, who view the wolf most negatively (Bisi
et al., 2010), represent a significant proportion of local peo-
ple in rural areas (Treves, 2009; Bisi et al., 2010; Pohja-
Mykr€a, 2016) and where effective guarding of wildlife is
practically impossible.
In this study, we suggest a quantitative method to include
the number of illegally killed wolves in the estimate of total
loss in advance, based on predicted poaching probabilities. A
concrete estimate of the number of illegally killed wolves
could diminish the acceptance of illegal killing through
group discipline if the number of hunting permits was bound
to poaching in a compensatory manner. A more stable local
wolf population, through the enhanced survival of breeding
adults, could be used as a measure of the absence of poach-
ing, which could lead to an increased number of legal per-
mits to hunt juveniles of local packs. The method of
predicted poaching probabilities is also applicable to other
species and ecosystems and can be used on different scales
that are appropriate for the system studied.
Local people are in key positions in anti-poaching actions
worldwide (Dickman, 2010; Challender & MacMillan, 2014;
Hazzah et al., 2017). Successful anti-poaching actions
require concrete estimates of poaching mortality and its rela-
tionship with legal harvesting. Engaging local hunters to sup-
press poaching to get more legal hunting permits could
stabilize the fluctuating wolf population by enhancing the
survival of breeding adults and ultimately could diminish all
aspects of wolf conflict.
Acknowledgments
We sincerely thank Sanna Kokko, Samuli Heikkinen, Seppo
Ronkainen, Markus Suominen, Leo Korhonen, Reima Ovaskai-
nen, Antero Hakala, and Esa Leinonen for their effort in data
collection. We thank H akan Sand and the two anonymous ref-
erees for providing valuable comments to the manuscript. Doc-
toral studies of J.Suutarinen were supported by the Finnish
Foundation for Nature Conservation and the Thule Institute’s
Aurora Doctoral Program, University of Oulu.
420
J. Suutarinen and I. Kojola
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