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Received: 25 June 2019 Revised: 17 September 2019 Accepted: 23 September 2019
DOI: 10.1111/conl.12679
LETTER
An experimental test of community-based strategies for
mitigating human–wildlife conflict around protected areas
Paola S. Branco M.S.1Jerod A. Merkle Ph.D.2Robert M. Pringle Ph.D.3
Lucy King Ph.D.4,5 Tosca Tindall6Marc Stalmans Ph.D.7Ryan A. Long Ph.D.1
1Department of Fish and Wildlife Sciences,
University of Idaho, Moscow, Idaho
2Department of Zoology and Physiology,
University of Wyoming, Laramie, Wyoming
3Department of Ecology and Evolutionary
Biology, Princeton University, Princeton,
New Jersey
4Elephants and Bees Project, Save the
Elephants, Nairobi, Kenya
5Department of Zoology, University of
Oxford, Oxford, UK
6Human Sciences Institute, University of
Oxford, Oxford, UK
7Department of Scientific Services,
Gorongosa National Park, Sofala,
Mozambique
Correspondence
Ryan Long and Paola Branco,Depar tment
of Fish and Wildlife Sciences, Universityof
Idaho, Moscow,ID 83844.
Email: ralong@uidaho.edu,
paola.medvet@gmail.com
Funding information
Save the Elephants; RuffordFoundation;
Campizondo Foundation; Artipopart; National
Science Foundation, Grant/AwardNumbers:
IOS-1656527, IOS-1656642
Abstract
Natural habitats are rapidly being converted to cultivated croplands, and crop-raiding
by wildlife threatens both wildlife conservation and human livelihoods worldwide.
We combined movement data from GPS-collared elephants with camera-trap data
and local reporting systems in a before–after-control-impact design to evaluate
community-based strategies for reducing crop raiding outside Mozambique’s Goron-
gosa National Park. All types of experimental fences tested (beehive, chili, beehive
and chili combined, and procedural controls) significantly reduced the number of
times elephants left the Park to raid crops. However, placing beehive fences at a subset
of key crossing locations reduced the odds that elephants would leave the Park by up
to 95% relative to unfenced crossings, and was the most effective strategy. Beehive
fences also created opportunities for income generation via honey production. Our
results provide experimental evidence that working with local communities to modify
both animal behavior and human attitudes can mitigate conflict at the human–wildlife
interface.
KEYWORDS
African savanna elephant, beehive fences, chili fences, crop raiding, human-dominated landscapes, key-
stone species, Loxodonta africana, movement corridors
1INTRODUCTION
The availability of high-quality forage in cultivated croplands
attracts wildlife (e.g., Middleton et al., 2017), and crop
raiding causes billions of dollars in economic losses every
year (Conover, 2002). Crop raiding by elephants (Loxodonta
africana,Elephas maximus) poses an especially severe
threat to human livelihoods in agroecosystems of Africa
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original
work is properly cited.
© 2019 The Authors. Conservation Letters published by Wiley Periodicals, Inc.
and Asia (Chiyo, Cochrane, Naughton, & Basuta, 2005;
O’Connell-Rodwell, Rodwell, Rice, & Hart, 2000; Shaffer,
Khadka, Van Den Hoek, & Naithani, 2019) and often occurs
along the boundaries of protected areas, where close prox-
imity of dense human and wildlife populations exacerbates
human–wildlife conflict (Bruner, Gullison, Rice, & da Fon-
seca, 2001; Wittemyer, Elsen, Bean, Burton, & Brashares,
2008).
Conservation Letters. 2020;13:e12679. wileyonlinelibrary.com/journal/conl 1of8
https://doi.org/10.1111/conl.12679
2of8 BRANCO ET AL.
Because human–elephant conflict involves both elephants
and humans, efforts to foster coexistence should ideally inte-
grate the modification of both elephant behavior (Mumby &
Plotnik, 2018) and human behavior/perceptions, the latter of
which are shaped by myriad factors (Dickman, 2010; Treves
& Bruskotter, 2014). Attitudes toward wildlife and protected
areas are influenced not only by crop losses per se, but also by
the degree to which individual beliefs and values are included
in decision-making processes (Infield, 2001; Bennet et al.,
2016). Thus, working closely with communities that are expe-
riencing conflict to foster relationships and establish rapport,
and then equipping them to participate directly in the mitiga-
tion process, may be an effective means of fostering human–
wildlife coexistence through a combination of decreased crop
losses and increased tolerance of elephants among community
members (Madden, 2004; Shaffer et al., 2019).
The coupling of animal deterrents with tangible incentives
to humans also holds powerful potential for fostering long-
term coexistence between humans and elephants. Indeed, off-
setting economic losses is considered essential to manag-
ing human–elephant conflict successfully (Hartter, Solomon,
Ryan, Jacobson, & Goldman, 2014; Snyman, 2014). Although
programs for compensating subsistence farmers for crop
losses to elephants have met with difficulties in Africa (Shaf-
fer et al., 2019), the production of marketable commodities
such as honey (King, Lala, Nzumu, Mwambingu, & Douglas-
Hamilton, 2017), chili products (Hedges & Gunaryadi, 2010),
or other cash crops (Parker & Osborn, 2006) as a byproduct of
deterrence can increase community buy-in and foster greater
tolerance toward elephants (Shaffer et al., 2019).
We studied human–elephant conflict along the southern
border of Gorongosa National Park, Mozambique, where the
elephant population is recovering from decimation by a civil
war that ended in 1992 (Pringle, 2017; Stalmans, Massad,
Peel, Tarnita, & Pringle, 2019). The goal of our project was
simultaneously to reduce the frequency of crop-raiding by ele-
phants and to improve attitudes toward elephants by work-
ing with community members to develop and test multiple
mitigation techniques with the potential to produce profitable
byproducts. We evaluated the efficacy of three techniques for
reducing elephant crop-raiding: (1) beehive fences (Karidozo
& Osborn, 2005; King, Lawrence, Douglas-Hamilton, & Voll-
rath, 2009; King, Douglas–Hamilton, & Vollrath, 2011; King
et al., 2017; Scheijen, Richards, Smit, Jones, & Nowak, 2018);
(2) chili-pepper fences (Hedges & Gunaryadi, 2010; Wiafe &
Sam, 2014); and (3) a combination of the two that we termed
“spicy beehive” fences. We conducted a manipulative experi-
ment in which we used two independent data streams (move-
ment data from GPS-collared elephants, and daily reports
from community members about the presence of elephants at
each fence location) to compare: (1) use of crossing points
by GPS-collared elephants between years with (year 2) and
without (year 1) fences; and (2) use of crossing points with
treatment, procedural-control, or no fences during year 2. We
hypothesized that fences of any kind would reduce the number
of times elephants exited the Park in year 2 (H1). We further
hypothesized that “spicy beehive” fences would be most effec-
tive for reducing crop raiding by elephants, followed by bee-
hive fences, chili fences, and procedural-control fences (H2).
2MATERIAL AND METHODS
2.1 Study area
In the Rift Valley of Gorongosa, annual precipitation averages
roughly 840 mm and occurs mostly between November and
March (Tinley, 1977). The Park is surrounded by a 5,333-km2
“buffer zone” where an estimated 200,000 people currently
reside. A large proportion of these residents cultivate crops
along the southern boundary of the Park, which is formed by
the Pungue River (Figure 1). From the 1970s–1990s, >90%
of Gorongosa’s 2,500+elephants were killed to feed soldiers
and to finance the purchase of arms during the Mozambican
Civil War (Convery & Morley, 2014; Stalmans, 2012). Ele-
phants are now recovering under the auspices of the Goron-
gosa Project (Pringle, 2017), and the most recent aerial census
counted roughly 600 individuals (Stalmans et al., 2019). After
the war, however, much of the buffer zone has been converted
to agricultural lands (Figure S1, Appendix S1), which strongly
attract elephants (Branco et al., 2018).
2.2 Animal capture and location data
We fit 12 adult male elephants (six in December 2015 and
six in August 2016), all of which were captured <1km
from crops, with GPS collars (Model AWT IM-SAT, Africa
Wildlife Tracking, Pretoria, South Africa) that were pro-
grammed to transmit a location every 30 minutes for 2 years
through the iridium satellite system. (Male elephants are gen-
erally more prone to crop-raiding behavior: Hoare, 1999.)
A detailed description of our capture and handling proce-
dures is provided by Branco et al. (2018); all procedures
were approved by the Animal Care and Use Committee at
the University of Idaho (protocol #2015–39). In addition, our
research was certified as exempt from continuing review by
the Institutional Review Board at the University of Idaho.
2.3 Community-based data
To evaluate the relative effectiveness of different fence types
at preventing elephants from crossing the Pungue to raid
crops (i.e., to test H2), we hired and trained a group of six
community members to work as project monitors during year
2 of the study. Each monitor was responsible for monitor-
ing 1–4 treatment or control fences (depending on distance
to their home and size of the fences), which they visited
BRANCO ET AL.3of8
FIGURE 1 Map of our study area in Gorongosa National Park, Mozambique in southeastern Africa. The Pungue River forms much of the
southern boundary of the Park, where we conducted our experiment. The Park is surrounded by a 5,333-km2buffer zone where ∼200,000 people
currently reside, many of whom are subsistence farmers
each morning to complete reports on whether elephants had
approached the fence, whether they had crossed the fence, and
the approximate number of elephants that had visited the loca-
tion (based on elephant footprints and fresh dung, damage to
fences, and photos from camera traps; Appendix S2). Mon-
itors were responsible for daily maintenance of fences and
camera traps during the mitigation experiment, and received
full-time salaries from Gorongosa National Park (commensu-
rate with those of full-time science technicians employed by
the park), as well as uniforms and bicycles to facilitate access
to their assigned areas in communities along the Pungue
River.
2.4 Randomized mitigation experiment
We used elephant GPS-collar data in combination with
information gleaned from community members to visually
identify locations where elephants routinely crossed the
Pungue to raid crops. We then visited all of the known cross-
ing locations that were used by elephants to access four of
the most-affected communities in the buffer zone—Micheu,
Madangua, Vinho, and Bebedo—which were dispersed along
∼18.7 km of the Pungue River (n=18 locations). We only
had sufficient resources to construct 13 fences as part of
our mitigation experiment, and thus we randomly selected
13 of these 18 crossings as treatment locations (Figure S3,
Appendix S3). Two of these locations could not be accessed
by vehicle and were therefore excluded from the study and
replaced by random selection from the five crossings that
were not selected as treatment locations (vehicular access
was essential for ensuring the safety of the research team
and for transporting fence construction materials to the
crossing locations). The remaining three accessible crossing
locations were left unmanipulated; there were no systematic
4of8 BRANCO ET AL.
FIGURE 2 Examples of elephant-deterrent strategies evaluated as part of our mitigation experiment in the buffer zone of Gorongosa National
Park (the park is to the left of each fence, farmlands to the right). (a) Chili fence. (b) Spicy beehive fence (beehive fences had the same design, but
hives were connected by baling twine). (c) Fake chili fence (control). (d) Fake beehive fence (control)
differences in river width, width of the crossing path, or
proximity to roads or agriculture between crossings where
fences were placed and crossings that remained unfenced.
For example, river width averaged 155 ±30.3 m (SE)at
locations where fences were constructed and 154 ±7.0 m
at unfenced locations. We assigned the four fence types
to the 13 treatment locations in a completely randomized
manner.
We constructed beehive fences (free-swinging hives con-
nected with light bailing twine) at three crossings, chili
fences (cotton fabric soaked in chili-impregnated vegetable
oil and interwoven with sisal ropes) at three crossings, and
spicy beehive fences (a combination of beehive and chili
fences) at three crossings. In addition, we constructed beehive
procedural-control fences (wooden planks of similar shape,
size, and color as active hives; Figure 2) at two crossings and
chili procedural-control fences (ropes without chili) at two
crossings. Detailed descriptions of fence design and construc-
tion are in Appendix S2.
We left fences in place for 3 months (September 17 to
December 20 2017) and evaluated results of the experiment
using two independent data streams: (1) movements of GPS-
collared elephants; and (2) daily reports from project moni-
tors, which included assessments of elephant sign (e.g., tracks
and dung), fence damage, and photos from camera traps. GPS-
collar data were collected throughout 2016–2017, and allowed
us to compare use of crossing locations before versus after
fences were erected (H1), and to evaluate the effectiveness
of different fence types at preventing elephants from crossing
the river (H2). Project monitors collected data only during the
mitigation experiment, and thus monitor data were used to test
H2 only.
2.5 Statistical analysis
We used generalized linear models (GLMs) to test our
hypotheses that (H1) fences of any type would reduce the
number of times elephants crossed the river to raid crops,
and that (H2) spicy beehive fences would be most effective
for reducing river crossings, followed by beehive fences, chili
fences, and procedural-control fences. For analyses of GPS-
collar data, the number of times collared elephants exited the
Park at each of the 16 crossing locations was used as the
response variable in a Poisson GLM. For analyses of mon-
itor data, the proportion of fence approaches by elephants
that ended in a river crossing (as opposed to a return to
BRANCO ET AL.5of8
the park) was used as the response variable in a binomial
GLM. Detailed descriptions of our statistical analyses are in
Appendix S4.
3RESULTS
In our focal stretch of ∼18.7 km of river adjacent to the
four heavily affected settlements, there were 67 crossings by
GPS-collared elephants between September 17 and Decem-
ber 20, 2016, but only 32 crossings during that same period
in 2017 after fences were erected (Table S5a, Appendix S5).
The mean number of crossings by GPS-collared elephants
at locations that remained unmanipulated (unfenced) during
the mitigation experiment doubled from 3.3 (95% CI =1.25–
5.35) in 2016 to 6.7 (95% CI =3.79–9.61) in 2017, suggesting
that elephants increased their use of unfenced crossings when
fences were erected at alternative crossing locations (note
that the majority of these posttreatment crossings occurred
at a single unfenced location, NF2; Table S5a, Appendix
S5). In contrast, the mean number of crossings at locations
with fences declined from 4.4 (95% CI =3.27–5.53) in
2016 (before fences) to 1.0 (95% CI =0.46–1.54) in 2017
(after fences) (Figure 3). The Treatment ×Year interaction
was highly significant (p<.001), consistent with our pre-
diction that there would be significantly fewer crossings at
fenced than at unfenced locations in 2017, after fences were
constructed. Park-wide (i.e., along the entire length of the
Pungue River that borders the Park), the total number of
crossings by GPS-collared elephants was 766 in 2016 (before
fences) and 744 in 2017 (after fences).
Camera-trap imagery indicated that elephants were,
in general, cautious in their interactions with fences (see
Appendix S6 for more detailed information on elephant
behavioral responses). All fence types, including procedural
controls, significantly (all p<.05) reduced the number of
times elephants crossed the river during the experiment
in 2017 (Figure 4). Mean (±95% CI) predicted number of
crossings per fenced crossing location (relative to unfenced
crossing locations) was lowest at locations with beehive
fences ( 𝑥 =2.99 ±2.01 fewer crossings), followed by spicy
beehive fences ( 𝑥 =1.90 ±1.21 fewer crossings), procedural-
control fences ( 𝑥 =1.67 ±0.98 fewer crossings), and chili
fences ( 𝑥 =1.61 ±1.07 fewer crossings; Figure 4; Table S5a,
Appendix S5), although none of the differences among fence
types were statistically significant after controlling for mul-
tiple comparisons. Odds ratios (i.e., exponentiated regression
coefficients) indicated that experimental fences reduced the
odds of an elephant crossing the river by anywhere from 80%
(chili fences) to 95% (beehive fences).
The data collected by project monitors during the exper-
iment told a story that was qualitatively similar to the data
from GPS-collared elephants. The mean (±95% CI) predicted
proportion of approaches by elephants that resulted in a river
FIGURE 3 Mean number of river crossings (±95% CI) by
GPS-collared elephants (n=12) at crossing locations that were or were
not blocked by fences during our mitigation experiment from
September 17 to December 20, in 2016 (prior to fence construction,
when all crossings remained unobstructed) and in 2017 (after fences
had been constructed at some crossings). The Treatment ×Year
interaction was significant (p<.001) in a Poisson GLM; p-values for
the main effects of Treatment (Fence vs. No fence) and Year were
p=.42 and p=.07, respectively
crossing was lowest (relative to procedural-control fences) at
beehive fences ( 𝑥 =25% ±35% fewer crossings), followed
by spicy beehive fences ( 𝑥 =7% ±35% fewer crossings)
and chili fences ( 𝑥 =25% ±35% more crossings; Figure 4;
Table S5b, Appendix S5). Again, however, none of the dif-
ferences among fence types were statistically significant after
controlling for multiple comparisons.
4DISCUSSION
All fence types in our study reduced crop-raiding excursions
by elephants, providing support for our hypothesis (H1)
that use of crossing points by elephants would decline in
year 2 after fences were constructed. This result suggests
that several different mitigation techniques can effectively
reduce crop-raiding; combining multiple techniques may also
help to minimize the potential for habituation (Hoare, 2015;
Shaffer et al., 2019). Prior research has demonstrated that
elephants quickly habituate to harmless mitigation methods
(O’Connell-Rodwell et al., 2000), and we suspect that the
effect of procedural-control fences in particular would quickly
attenuate. On the contrary, honey bees are naturally avoided
by elephants (King et al., 2011), and thus the likelihood of
habituation is largely dependent upon hive occupation (King
et al., 2017). Beehive fences were especially effective in our
study, and hive occupancy reached 94% in the first 7 weeks
following fence construction, likely aided by the proximity
6of8 BRANCO ET AL.
FIGURE 4 Parameter estimates (with 95% CI) from binomial
GLMs of (A) the number of river crossings by GPS-collared elephants
during our mitigation experiment (September 17 to December 20,
2017), and (B) the proportion of fence approaches by elephants that
resulted in a river crossing during that same period (as determined by
track, dung, and camera-trap data collected by project monitors).
Parameter estimates in (A) indicate the predicted number of river
crossings per crossing location at treatment sites with each fence type,
relative to unfenced locations (all p<.05), whereas parameter
estimates in (B) indicate the predicted proportion of fence approaches
that resulted in a river crossing at locations with each treatment fence
type, relative to control fences that lacked the putative deterrence
mechanisms (i.e., chilies and/or bees; all p>.15)
of a stable water source (i.e., the Pungue River) (King et al.,
2017). These results are similar to those of previous studies
that evaluated the use of beehive fences (e.g., King et al.,
2009; King et al., 2011; King et al., 2017; Scheijen et al.,
2018) or chili fences (e.g., Chang’a et al., 2016; Gunaryadi,
Sugiyo, & Hedges, 2017) around individual farms. Our work
builds on those studies by demonstrating the efficacy of
using discontinuous fencing to block key corridors used by
elephants to access crops rather than fencing individual farms
or erecting continuous fences along entire protected-area
boundaries (at great expense). Our results suggest that if com-
munities can tolerate some (much-reduced) level of crop raid-
ing, then discontinuous beehive fencing at crossing sites may
represent a compromise solution that simultaneously reduces
both the incidence of crop raiding and the costs of mitigation.
Another important benefit of beehive fences is their poten-
tial to generate revenue from honey production. Gorongosa
currently supports an apiculture program in the buffer zone,
and we constructed our beehive fences using the same hives
as those used by Park apiculturists. Park honey producers can
harvest 10–14 kg of honey per hive annually, which they can
sell for 60–70 Meticais ($1–1.5 USD) per kg. Thus, a sin-
gle beehive fence consisting of 15 hives could produce 150–
210 kg of honey per year, generating $150–315 USD. This is
2–4 times the current minimum annual wage in Mozambique.
By way of comparison, construction of a beehive fence with
15 hives in our study cost ∼$773 USD in materials. The hives
themselves comprised the majority of the cost ($33.50 USD
apiece for Kenyan top bar hives), with other equipment and
supplies (bee attractant, hardwood poles, yellow paint, bail-
ing twine, nails, wire, bee brush, and gloves) totaling ∼$270
USD. A detailed list of costs associated with construction of
all fence types is provided in Appendix S7 (Table S7). At
Gorongosa, the initial cost of constructing fences for our study
was borne by the park and various donors, and community
members were responsible for subsequent maintenance, as
well as the harvest and sale of honey. Such collaborative cost-
sharing arrangements hold potential for fostering coexistence
between humans and elephants by simultaneously reducing
the frequency of crop raiding, by demonstrating the commit-
ment of protected areas to both human and elephant well-
being, and by providing economic incentives to community
members through both reduced crop losses and the sale of
honey.
Spicy beehive fences did not repel elephants as consistently
as beehive fences alone. Although these results are contrary to
our hypothesis (H2), camera-trap imagery suggests that they
may have stemmed from differences in weight of the materials
used to link hives together in beehive fences versus spicy bee-
hive fences. The interwoven cotton strips and sisal ropes used
to sustain the chili mixture (see Appendix S2) were consid-
erably heavier than the simple twine ropes used to construct
the beehive fences (Figure 2). As a result, these ropes sagged
more, making it easier for elephants to step over them. With
the chili fences, it was possible to tie the ropes very tightly to
the bamboo poles. In contrast, with the spicy beehive fences
it was necessary to leave the chili ropes looser to keep from
pulling the hives sideways. Future work to improve the design
of this strategy by keeping ropes at a height that prevents
elephants from stepping over them would likely improve its
effectiveness.
Our experimental design required leaving some key cross-
ing points unfenced, and elephants increased their use of those
crossings following fence construction (Figure 3). Osipova
et al. (2018) reported similar results following construction
BRANCO ET AL.7of8
of electrified fences in the Amboseli ecosystem in Kenya.
Despite increased use of unfenced crossings by elephants dur-
ing our experiment, however, the total number of crossings
that occurred in our study area was reduced by more than half
following construction of fences. This suggests that fences
used in our study—and beehive fences in particular—shifted a
considerable amount of elephant activity to other stretches of
the river even though some crossing locations remained unob-
structed. This shift in elephant activity away from commu-
nities that participated in our study likely contributed to the
generally positive perceptions of our project among commu-
nity members (Appendix S8). It is not possible to know from
our data how fencing all crossing points might have influ-
enced elephant crop-raiding behavior in our study area. How-
ever, based on results of our study, the Conservation Depart-
ment at Gorongosa recently began large-scale deployment
of beehive fences at crossings all along the Pungue River.
This management experiment will provide opportunities to
test the consequences of full-scale implementation of beehive
fences.
Protected areas form the cornerstone of efforts to conserve
biodiversity worldwide (Bruner et al., 2001; Hilborn et al.,
2006). Crop raiding by wildlife undermines the effectiveness
of protected areas—which is already tenuous owing to fund-
ing shortfalls (Lindsey et al., 2018)—by bringing wildlife into
direct conflict with human populations. The future sustain-
ability of many protected areas will depend in part on devel-
oping strategies for mitigating human–wildlife conflict that:
(1) are affordable to implement; (2) can be maintained by
local communities; (3) incentivize communities by providing
avenues of economic gain (e.g., reduced crop losses coupled
with revenue from honey production); and (4) help to alter
perceptions of wildlife among community members (Shaffer
et al., 2019). Our study experimentally demonstrates the
potential effectiveness of one such strategy, and our approach
can be further refined, adapted, and scaled up to reduce crop
raiding by megaherbivores and other wildlife around pro-
tected areas.
ACKNOWLEDGMENTS
This work and its participants were supported by the
U.S. National Science Foundation (IOS-1656642 to R.A.L.;
IOS-1656527 to R.M.P.). We also thank the University
of Idaho, Gorongosa Project, Rufford Foundation, Campi-
zondo Foundation, Artipopart, Save the Elephants, Sr. Jean-
Marc Grün and Victoria Branco for the financial support.
We are grateful to our field assistants Castiano Lencas-
tro, Michel Souza and Elyce Gosselin. We also thank Janet
Rachlow, Joyce Poole and Petter Granli for their comments
and support. The data that support the findings of our
study are publicly available in the Dryad Digital Repository
(doi 10.5061/dryad.m63xsj3xd).
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SUPPORTING INFORMATION
Additional supporting information may be found online in the
Supporting Information section at the end of the article.
How to cite this article: Branco PS, Merkle JA,
Pringle RM, et al. An experimental test of community-
based strategies for mitigating human–wildlife con-
flict around protected areas. Conservation Letters.
2020;13:e12679. https://doi.org/10.1111/conl.12679
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