Scientific RepoRts | 7:45222 | DOI: 10.1038/srep45222
Hunting promotes spatial
reorganization and sexually
M. Leclerc1, S. C. Frank2, A. Zedrosser2,3, J. E. Swenson4,5 & F. Pelletier1
Harvest can aect the ecology and evolution of wild species. The removal of key individuals, such as
matriarchs or dominant males, can disrupt social structure and exacerbate the impact of hunting on
population growth. We do not know, however, how and when the spatiotemporal reorganization takes
place after removal and if such changes can be the mechanism that explain a decrease in population
growth. Detailed behavioral information from individually monitored brown bears, in a population
where hunting increases sexually selected infanticide, revealed that adult males increased their use
of home ranges of hunter-killed neighbors in the second year after their death. Use of a hunter-killed
male’s home range was inuenced by the survivor’s as well as the hunter-killed male’s age, population
density, and hunting intensity. Our results emphasize that hunting can have long-term indirect eects
which can aect population viability.
Human activities are a major evolutionary force aecting wild populations1. ere is increasing evidence that
human exploitation leads to changes in morphological and life history traits worldwide1–4. For example, recent
studies have shown that size-selective harvest by commercial sheries and trophy hunting can induce evolution of
heritable traits5–9. Harvest-induced evolution might not be desirable as the selection induced by human exploita-
tion can be in the opposite direction of natural selection10–12.
Hunting can also have indirect eects on wildlife, although such eects are oen ignored by managers, even
though the removal of key individuals by hunting could change a population’s social structure13. For example,
simulations suggest that the social networks of killer whales (Orcinus orca) may be vulnerable to targeted removal
of individuals14. In African elephants (Loxodonta africana) the enhanced discriminatory abilities of the oldest
individuals inuences the social knowledge and reproductive success of entire groups15, suggesting that the loss
of older individuals could decrease the tness of all females within the group. In social species, the removal of
any individual could aect social dynamics by changing the social structure. However, empirical evidence link-
ing hunting and spatiotemporal reorganization of the social structure is lacking and the data needed to investi-
gate this question are rarely available. Given the large number of species targeted by harvest, understanding the
potential eects of removal on subsequent space use, social structure, and the tness consequences for surviving
individuals is critical to achieve sustainable hunting practices.
Here, we used detailed individual behavioral information from a Scandinavian brown bear (Ursus arctos) pop-
ulation (monitored from 2008–2015) to evaluate whether surviving adult males (hereaer referred to as survivors)
shi their home range use aer a neighboring adult male has been killed by hunting (TableS1). We further investi-
gated the intrinsic and extrinsic factors driving the spatiotemporal reorganization of male spatial structure. In this
population, the removal of adult males through hunting increases the risk of sexually selected infanticide (SSI)16,17,
which is a major determinant of population growth18. Although important for sustainable wildlife manage-
ment19, the mechanism behind the harvest-induced increase of SSI remains unknown [but see Loveridge et al.20].
1Canada Research Chair in Evolutionary Demography and Conservation & Centre for Northern Studies, Département
de biologie, Université de Sherbrooke, Sherbrooke, J1K2R1, Canada. 2Faculty of Technology, Natural Sciences, and
Maritime Sciences, Department of Natural Sciences and Environmental Health, University College of Southeast
Norway, N-3800 Bø i Telemark, Norway. 3Department of Integrative Biology, Institute of Wildlife Biology and Game
Management, University of Natural Resources and Life Sciences, Vienna, Gregor Mendel Str. 33, A - 1180 Vienna,
Austria. 4Faculty of Environmental Sciences and Natural Resource Management, Norwegian University of Life
Sciences, PO Box 5003, NO - 1432 Ås, Norway. 5Norwegian Institute for Nature Research, NO-7485 Trondheim,
Norway. Correspondence and requests for materials should be addressed to M.L. (email: Martin.Leclerc2@
Received: 01 December 2016
Accepted: 21 February 2017
Published: 23 March 2017
Scientific RepoRts | 7:45222 | DOI: 10.1038/srep45222
Spatial reorganization due to hunting of males may be the responsible mechanism, by increasing the probability
that a female will encounter a new male that is unlikely to be the father of her cubs13,16.
We found that survivors increased their use of the home ranges of hunter-killed males in the second year aer
their death (Fig.1, TableS2). is time lag in the response likely is related to the bear’s ecology. Bears den from
October to April21,22, shortly aer the hunting season in late August—September. e size of the annual home
range in our study population is mainly dened by space use during the mating season (May to mid-July), when
males exhibit a roam-to-mate behavior23. erefore, we hypothesize that survivors do not readjust their home
range until aer the rst mating season without the hunter-killed neighbor. is could explain the two-year time
lag in spatial reorganization. Our results support the contention that the spatiotemporal reorganization of male
home ranges is an important mechanism linking hunter harvest to an increase in SSI, described above. It is also
consistent with earlier studies in the same population showing lower cub survival following a two-year time lag
aer a male had been killed16,17.
We further investigated which intrinsic (ages of hunter-killed and surviving males) and extrinsic factors (pop-
ulation density and hunting intensity) modulated the speed and strength of the survivors’ response to hunting
removals (Fig.2, TablesS3 and S4). e use of a hunter-killed male’s home range by its surviving neighbors
was inuenced by (in order of decreasing relative importance) survivor’s age (∆ BIC = 115), hunting intensity
(∆ BIC = 76), population density (∆ BIC = 74), and hunter-killed male’s age (∆ BIC = 6). Older survivors used
a hunter-killed male’s home range less strongly following the hunter-killed male’s death than younger survivors
(Fig.2A). is suggests that older males may already have held home ranges with better resources, including food
and females. Age-dependent home range quality could also explain why survivors increased their use of an old
hunter-killed male’s home range more than that of a younger hunter-killed male (Fig.2D).
Survivors more strongly increased their use of a hunter-killed male’s home range in the second year aer its
death when hunting intensity was greater (Fig.2B). As increasing hunting intensity will increase the number
of openings for surviving males, this should lead to a higher degree of spatial reorganization. We previously
reported that the killing of an adult male within 25 km of a female strongly reduced the survival of her cubs, with
a two-year time lag, although an increase in the number of killed males within 25 km had no signicant additive
eect17. Even though the degree of spatial reorganization increased with increased hunting intensity, this might
not always translate into a correspondingly lower cub survival, because even though more surviving males may
respond to increased hunting removal, only one infanticidal male is sucient to kill most of females’ cubs. e
other extrinsic factor aecting shis in a survivor’s home range use was population density (Fig.2C). Survivors
at higher densities had higher initial overlap with the hunter-killed male and showed a weaker reorganization
response than survivors at lower densities (Fig.2C). Stronger competition for space between neighbors might
explain why we observed higher initial overlap, with a weaker response at higher densities.
We identied a key behavioral mechanism linking hunting to an increase in SSI and show how post-hunt spa-
tiotemporal reorganization of males was modulated by both intrinsic and extrinsic factors. By removing males
from the population, hunters destabilized the spatial organization of the population for at least two years aer a
male had been killed. is period of two years might be specic to brown bears, due to their denning period and
could be dierent in other harvested species with SSI, such as lions (Panthera leo)20 or cougars (Puma concolor)24.
Nevertheless, hunting increases shis in home range use by surviving males and increases the probability of
SSI16,17. Male bears seem to assess their paternity through their mating history25, and increasing the magnitude
of shis in home range use would increase the probability that a male could encounter a female with whom he
had not previously mated. Such a pattern is expected regardless of the cause of death (e.g., vehicle collision, man-
agement kill, natural mortality). However, hunting is oen additive to natural mortality, as in our study system26,
which increases the occurrence of SSI compared to unharvested systems.
e spatial distribution of the hunting mortality of bears was not homogenous in our study area27. Spatial
and social relationships of bears are likely to change more rapidly in areas with higher hunting mortality, thereby
potentially decreasing the cohesion of their social network28,29 but see ref. 30. Such eects could also inuence
Figure 1. Changes in surviving male brown bears use of hunter-killed neighboring males’ home ranges
over time. Shown are the coecients and 95% condence intervals for three consecutive years, i.e. the year the
hunter-killed male was shot (baseline) and the following two years.
Scientific RepoRts | 7:45222 | DOI: 10.1038/srep45222
the female reproductive rate because female brown bears exhibit kin-related spatial structures31, where neighbors
negatively aect each other’s probability of having cubs32,33. e direct eect of removals due to hunting, in addi-
tion to the indirect eects of increasing cub mortality due to SSI and the potential impacts of decreasing social
network cohesion, all increases heterogeneity in survival and reproductive rates. ese eects combined could
increase demographic variability and ultimately aect eective population size34,35. erefore, we expect spatially
structured demographic variability that could potentially result in source-sink dynamics35,36.
Our study sheds light on the importance of animal behavior to explain time lags in the responses to hunting
in the wild. Understanding the indirect consequence of hunting over long time scales is critical for developing
sustainable management practices and for the viability of harvested populations.
e study area was in south-central Sweden (61°N, 15°E) and was composed of bogs, lakes, and intensively man-
aged coniferous forest stands. e dominant tree species were Norway spruce (Picea abies), Scots pine (Pinus
sylvestris), lodgepole pine (Pinus contorta), and birch (Betula spp.). Elevations ranged between 150 and 725 m asl.
Gravel roads (0.7 km/km2) were more abundant than paved roads (0.14 km/km2). See Martin et al.37 for further
information about the study area.
We captured brown bears from a helicopter using a remote drug delivery system (Dan-Inject® ,
Børkop, Denmark). We determined sex at capture and extracted a tooth from unknown individuals for age
Figure 2. Inuence of intrinsic and extrinsic factors on the speed and strength at which a surviving male
will use hunter-killed neighboring males’ home ranges. Shown are the coecients and 95% condence
intervals for three consecutive years, i.e. the year the hunter-killed male was shot (baseline) and the following
two years, depending on the surviving male’s age (A), hunting intensity (B), population density (C), and hunter-
killed male’s age (D), e low and high values in each panel represent the 25th and 75th percentiles, respectively,
observed in the database.
Scientific RepoRts | 7:45222 | DOI: 10.1038/srep45222
determination38. We equipped bears with GPS collars (GPS Plus; Vectronic Aerospace GmbH, Berlin, Germany)
programed to relocate a bear with varying schedules (≤ 1 hour intervals). See Fahlman et al.39 for details on
capture and handling. All captured bears were part of the Scandinavian Brown Bear Research Project and all
experiments, captures and handling were performed in accordance with relevant guidelines and regulations and
were approved by the appropriate authority and ethical committee (Naturvårdsverket and Djuretiska nämden i
Spatial analysis. We used adult male bears ≥ 4 years in the analysis to exclude natal dispersers40. We did
not include natal dispersers because all male dispersers moved outside the study area where too few or no other
males were GPS-collared. In addition, females actively defend their cubs during infanticide attempts. erefore,
younger dispersing males that have not yet attain full body size are less likely to successfully commit SSI than
older, larger and better established males41. We screened the relocation data of adult males and removed GPS xes
with dilution of precision values > 10 to increase spatial accuracy. To reduce autocorrelation, we used a 6-hour
minimum interval between successive positions for a given bear. We excluded bears in years for which an individ-
ual had < 75% of days with GPS locations from 1 May to 30 September.
We used an approach adapted from resource selection functions [RSFs;42] developed by Bischof et al.43. For
each GPS-collared hunter-killed male we (1) determined its annual 95% kernel home range for the active period
(1 May to 30 September or the day before he was killed) of the year in which he was killed and (2) calculated a
40-km radius circular buer centered on its home range centroid. is radius was used because it represents the
distance within which 95% of home range centroids of successful mates occurred44 and the distance at which
the eect of male removal on cub survival seems to disappear17. In a given year, we used GPS relocations of the
hunter-killed male and all the GPS locations of surviving adult males within the buer (hereaer called survivors)
to (3) calculate a 95% kernel isocline (hereaer called sampling space). For each survivor, we (4) generated as
many random than GPS relocations within the sampling space and (5) determined if GPS and random relocations
were inside or outside the hunter-killed bear’s home range. We repeated steps 3–5 for 3 consecutive years, i.e. the
year a hunter-killed male had been killed and the two following years. We updated the sampling space annually
by keeping the hunter-killed males’ relocations the year he was killed constant for the three years, and used the
appropriate relocations of survivors for each year. We only used survivors that were alive and monitored during
the three-year period. We repeated these steps for each hunter-killed male. is enabled us to test whether survi-
vors increased their use of a hunter-killed male’s home range the years following its death.
For each hunter-killed male we also extracted a population density index derived from county-level scat col-
lections in Sweden. We used the method of Jerina et al.45 and summed the weighted values of an individual bear’s
multiple scats across a grid of 10 × 10 km. is was carried out for each county separately, aer which the distri-
bution was corrected temporally, using county-level trends of the Large Carnivore Observation Index46,47, pro-
vided by the Swedish Association for Hunting and Wildlife Management. Lastly, we calculated a proxy of hunting
intensity based on the number of dead adult males located within the 40-km radius circular buer centered on
a given hunter-killed male’s home range centroid over a 3-year period prior to its death [see Gosselin et al.17 for
Statistical analysis. As a first step, we determined if surviving males shifted their home range use in
response to the removal of a hunter-killed male. To do so, we used a generalized linear mixed model (GLMM)
with binomial distributed errors. We coded the dependent variable either as GPS (coded 1) or random (coded
0) relocation. As independent variables we used a dummy variable representing whether the relocations were
inside (coded 1) or outside (coded 0) the hunter-killed males home range, as well as a variable representing the
period of the relocations (3-level factor; the year of the hunter-killed male’s death, as well as 1 and 2 years aer
the hunter-killed male’s death). We evaluated 4 candidate models (TableS1) and selected the most parsimonious
based on the Bayesian information criterion (BIC)48. To control for the eect of year and unequal sample sizes
across individuals, we included Year and the survivor ID nested within the hunter-killed males’ ID as random
intercepts in all candidate models.
In a second step, we examined how intrinsic (i.e., age of survivor and hunter-killed males) and extrinsic (i.e.,
population density and hunting intensity) factors inuenced the speed and strength at which a survivor would
adjust its home range use in response to the removal of a hunter-killed male. We used a GLMM with binomial
distributed errors and coded the dependent variable either as GPS (coded 1) or random (coded 0) relocation. We
evaluated the eect of six independent variables; inside vs outside the hunter-killed male home range, period,
age of the survivor, age of the hunter-killed male, population density, and hunting intensity to build 17 candidate
models (TableS3). We selected the most parsimonious model based on BIC. To control for the eect of year and
unequal sample sizes across individuals, we included Year and the survivor ID nested within the hunter-killed
males’ ID as random intercepts in all candidate models. To facilitate model convergence, we scaled (mean = 0,
variance = 1) all numerical covariates. We assessed the relative importance of variables within the most parsimo-
nious model by dropping each variable and monitoring the ∆ BIC. e larger the relative dierence in BIC com-
pared to the most parsimonious model, the more important we considered a variable. For all candidate models
tested, the variance ination factor (VIF) value was < 249. We used R version 3.2.3 for all statistical analyses50.
We captured and GPS-monitored a total of 15 adult males between 2008 and 2015. e database contained
19,133 GPS and 19,133 random relocations of 11 hunter-killed males and 7 survivors, for a total of 23 survivor –
hunter-killed male pairs.
Scientific RepoRts | 7:45222 | DOI: 10.1038/srep45222
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We would like to thank M. Festa-Bianchet, D. Garant, J. Van de Walle, one anonymous reviewer and the editor
for useful comments on earlier version of the manuscript. ML was supported nancially by NSERC and FRQNT.
FP was funded by NSERC discovery grant and by the Canada research Chair in Evolutionary Demography
and Conservation. is is scientic publication No. 232 from the SBBRP, which was funded by the Swedish
Environmental Protection Agency, the Norwegian Directorate for Nature Management, the Research Council
of Norway, the Austrian Science Fund, and the Swedish Association for Hunting and Wildlife Management.
We acknowledge the support of the Center for Advanced Study in Oslo, Norway, that funded and hosted our
research project “Climate eects on harvested large mammal populations” during the academic year of 2015–
2016 and funding from the Polish-Norwegian Research Program operated by the National Center for Research
and Development under the Norwegian Financial Mechanism 2009–2014 in the frame of Project Contract No
All authors participated in the study design. M.L. and S.C.F. carried out statistical analyses, F.P., J.E.S. and
A.Z. secured funding, J.E.S. and A.Z. coordinated the Scandinavian Brown Bear Research Project. All authors
participated in writing the manuscript.
Supplementary information accompanies this paper at http://www.nature.com/srep
Competing Interests: e authors declare no competing nancial interests.
How to cite this article: Leclerc, M. et al. Hunting promotes spatial reorganization and sexually selected
infanticide. Sci. Rep. 7, 45222; doi: 10.1038/srep45222 (2017).
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