Consequences of recreational hunting for biodiversity conservation and livelihoods

Abstract

The widespread activity of recreational hunting is proposed as a means of conserving nature and supporting livelihoods. However, recreational hunting-especially trophy hunting-has come under increasing scrutiny based on ethical concerns and the arguments that it can threaten species and fail to contribute meaningfully to local livelihoods. We provide an overview of the peer-reviewed literature on recreational hunting of terrestrial birds and mammals between 1953 and 2020 (>1,000 papers). The most-studied species are large mammals from North America, Europe, and Africa. While there is extensive research on spe-cies' ecology to inform sustainable hunting practices, there is comparably little research on the role of local perceptions and institutions in determining socioeconomic and conservation outcomes. Evidence is lacking to answer the pressing questions of where and how hunting contributes to just and sustainable conservation efforts. We outline an agenda to build this evidence base through research that recognizes diverse social-ecological contexts.

Full text

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Consequences of recreational hunting
for biodiversity conservation and livelihoods
Enrico Di Minin,
1,2,3,9,
*Hayley S. Clements,
1,4,9,
*Ricardo A. Correia,
1,2,5
Gonzalo Corte
´s-Capano,
1,2
Christoph Fink,
1,2
Anna Haukka,
1,6
Anna Hausmann,
1,2
Ritwik Kulkarni,
1,2
and Corey J.A. Bradshaw
7,8
1
Helsinki Lab of Interdisciplinary Conservation Science, Department of Geosciences and Geography, University of Helsinki, 00014 Helsinki,
Finland
2
Helsinki Institute of Sustainability Science, University of Helsinki, 00014 Helsinki, Finland
3
School of Life Sciences, University of KwaZulu-Natal, 4041 Durban, South Africa
4
Centre for Complex Systems in Transition, Stellenbosch University, 7600 Stellenbosch, South Africa
5
DBIO & CESAM – Centre for Environmental and Marine Studies, University of Aveiro, 3810-193 Aveiro, Portugal
6
Finnish Museum of Natural History, University of Helsinki, 00014 Helsinki, Finland
7
Global Ecology, College of Science and Engineering, Flinders University, Adelaide, South Australia 5001, Australia
8
ARC Centre of Excellence for Australian Biodiversity and Heritage, Adelaide, South Australia 5001, Australia
9
These authors contributed equally
*Correspondence: enrico.di.minin@helsinki.fi (E.D.M.), hayley.clements@helsinki.fi (H.S.C.)
https://doi.org/10.1016/j.oneear.2021.01.014
SUMMARY
The widespread activity of recreational hunting is proposed as a means of conserving nature and support-
ing livelihoods. However, recreational hunting—especially trophy hunting—has come under increasing
scrutiny based on ethical concerns and the arguments that it can threaten species and fail to contribute
meaningfully to local livelihoods. We provide an overview of the peer-reviewed literature on recreational
hunting of terrestrial birds and mammals between 1953 and 2020 (>1,000 papers). The most-studied spe-
cies are large mammals from North America, Europe, and Africa. While there is extensive research on spe-
cies’ ecology to inform sustainable hunting practices, there is comparably little research on the role of local
perceptions and institutions in determining socioeconomic and conservation outcomes. Evidence is lack-
ing to answer the pressing questions of where and how hunting contributes to just and sustainable conser-
vation efforts. We outline an agenda to build this evidence base through research that recognizes diverse
social-ecological contexts.
INTRODUCTION
Human activities are eroding biodiversity and are consequently
reducing the benefits provided by nature to people.
1
Unsustain-
able harvesting and land-use change are historically the most
influential drivers of biodiversity loss.
2
When unsustainable, har-
vesting (e.g., hunting or fishing, see Salafsky et al.
3
for a
classification of subthreats within the topic of ‘‘unsustainable
harvesting’’) of species can lead to population declines and, ul-
timately, to extinction.
4
Infamous examples include the great
auk Pinguinus impennis, Steller’s sea cow Hydrodamalis gigas,
the passenger pigeon Ectopistes migratorius, the marsupial ‘‘ti-
ger’’ Thylacinus cynocephalus, and the dodo Raphus cucullatus.
Conservation actions in response to unsustainable harvesting
are multiple and can be multifaceted.
5
Strict protection (i.e., no
harvesting permitted) or legislated restrictions on harvesting
can be appropriate actions in response to unsustainable rates
of harvest, but these can prevent people who live alongside
biodiversity from benefiting from nature conservation.
6
A lack
of benefit to local people, particularly where it negatively impacts
their well-being, is both a problem of inequity and a conservation
issue when it results in increased wildlife poaching or persecu-
tion.
7
Hence, there is need to apply actions that generate posi-
tive benefits for conservation and local people, particularly in
places where pressure on species from unsustainable harvest-
ing is especially severe
8
and poverty is high.
9
Recreational hunting has long been promoted as a means of
generating benefits to conserve and restore biodiversity and
contribute to human well-being.
10,11
Proponents of recreational
hunting argue that the finances generated can prevent natural
habitat conversion and biodiversity loss and help support con-
servation actions (e.g., anti-poaching measures).
12
For example,
recreational hunting in the United States generates funding for
state and provincial conservation agencies.
13
In some countries
in sub-Saharan Africa,
11
recreational hunting areas cover more
land than formally protected areas.
14
In addition, recreational
hunting in some contexts provides livelihoods and other benefits
(e.g., meat, increased stakeholder collaboration) to local peo-
ple.
11,15
It is therefore often presented as a means of achieving
objectives for both biodiversity conservation and sustainable
development.
11
Other conservation benefits of recreational
hunting include, inter alia,
16
controlling populations of overabun-
dant species
17
and restoring ecosystems
18
and species’ popu-
lations.
19,20
Many concerns have, however, been levelled against the role
of recreational hunting in supporting biodiversity conservation
and livelihoods. One main concern is the sustainability of recre-
ational hunting in the face of increasing human pressures on
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biodiversity.
21
There is also uneasiness about the introduction of
alien species for hunting purposes and their impacts on ecosys-
tems.
22
Furthermore, the revenue generated from recreational
hunting does not always benefit local people living with the biodi-
versity in question.
11
Increasingly, the practice is considered
ethically and morally unjustifiable regardless of any positive out-
comes for biodiversity conservation.
23,24
In particular, many
consider trophy hunting (i.e., where the hunter retains some or
all of the animal’s body as a ‘‘trophy’’)
25
to be problematic
from an ethical perspective. There is increasing public opposi-
tion to the ethical legitimacy of relatively wealthy foreigners hunt-
ing large, charismatic mammals such as lions Panthera leo and
elephants Loxodonta africana.
24,26
Some also consider trophy
hunting to be unacceptable because it perpetuates the cultural
narrative of chauvinism, colonialism, and anthropocentrism.
25
There is therefore a growing movement to influence legislation
and ban imports of trophies.
27
Overall, debates centered on rec-
reational hunting are among the most polarizing in conservation
science and practice today.
28
Here, we provide an overview of the peer-reviewed literature
on recreational hunting globally. Specifically, we use topic
modeling to investigate and summarize the main topics exam-
ined and the geographic and taxonomic focus in recreational-
hunting research. Drawing on these topics, we consider the
diverse implications of recreational hunting for nature conserva-
tion and the livelihoods and well-being of people. We conclude
by outlining a research agenda to provide evidence-based rec-
ommendations for enabling the intersection of sustainability, eq-
uity, and ethics when designing best-practice hunting man-
agement.
WHAT IS RECREATIONAL HUNTING?
Hunting can be divided into three broad categories: ‘‘subsis-
tence hunting’’ (food for own consumption), ‘‘commercial hunt-
ing’’ (sale of animal products to a consumer community in local,
national, or international markets), and ‘‘recreational hunting.’’
29
Recreational hunting can be broadly defined as the pursuit and
killing of animals primarily for leisure and enjoyment purposes.
30
However, there may be additional motivations besides recrea-
tion, including meat and trophy acquisition, spiritual, social,
and cultural motivations, added exercise, the physiological ef-
fects of excitement, and the desire to make a direct or indirect
contribution to population management, invasive alien species
control, and habitat conservation.
31–33
For example, hunting
the fox Vulpes vulpes in the United Kingdom was largely consid-
ered to be a socially and culturally meaningful recreational activ-
ity but also a means to control fox populations.
34
Equally, many
deer hunters pursue trophies, but also hunt for the experience
and for food. Such complexities of motivations can prevent clear
distinction between recreational, subsistence, and commercial
hunting.
30
A range of species, particularly mammals and birds, are tar-
geted by recreational hunters who pursue these species by using
a range of methods, including bows and arrows, dogs, and
guns.
32
Recreational hunting includes both local hunting and
tourism hunting.
32
Local recreational hunting is done by hunters
that live near the hunting area. Recreational-hunting tourism is
done by hunters that travel longer distances from home, often
abroad, and who are prepared to pay considerably for the oppor-
tunity of hunting particular species.
Recreational hunting is often regulated, with the intention of
meeting certain ecological, social, ethical, and sustainability
standards (at least in theory).
35
Regulation normally occurs via
national or subnational legislation, which recognizes the role of
specific authorities for management. Legislation might also be
accompanied by local to regional by-laws regulating recreational
hunting, and in some instances by local traditional institutions.
International regulations also play an important role in the
cross-border movements of hunting trophies from certain spe-
cies. Legislating agencies normally determine the methods for
regulating the numbers of animals hunted and/or seasons in
which to hunt. Sustainability can be promoted by establishing
a fixed hunting quota, by regulating hunting effort temporally
and/or spatially, or by limiting the proportion of the population
that can be hunted. A review of such regulations goes beyond
the scope of our study, and the reader is referred to Rosser
35
for a detailed discussion.
ETHICS OF RECREATIONAL HUNTING
Recreational hunting is a controversial and contested issue. The
public acceptance of any form of recreational hunting, even in
relation to invasive alien species,
36
is today lower than it has
ever been in the past,
37
with concerns regarding animal welfare
and rights dominating the discussion and ethical apprehension
on the morality of hunting for pleasure.
38
The ethical arguments
against hunting can originate from the perspective of animal wel-
fare focusing on the rights of the individual animal and the moral-
ity of eating meat, which contrasts with the land ethic justifying
population management via hunting for the greater good of the
ecosystem rather than the individual animal.
39
Another ethical
debate emerges between primitivist arguments of hunting as
an evolved human behavior that implies ‘‘trophic responsibil-
ity,’’
40
and the ecofeminist argument that primitivism marginal-
izes women’s relationship with nature and that recreational
hunting is the outcome of the patriarchy’s fixation on violence
and death.
39
The common justification for recreational hunting
as an experiential fulfillment has been argued to be morally
inconsistent, because its protagonists create a false hunting ide-
ology given that commoditization of the prey weakens justifica-
tion regarding the thrill of the chase.
41
Furthermore, there are
philosophical arguments that expose the fallacy of attempting
to distinguish the morality of sport versus subsistence hunting,
and that ethical hunting of any type cannot rely on human con-
structs and should be instead couched in terms of ecology and
evolution.
42
Trophy hunting in particular is a charged emotional stage, with
entire treatises devoted to understanding why elements of soci-
ety are either strongly supportive of or morally outraged by the
idea.
43
Indeed, much of the incentive can be traced to its colonial
heritage as an elitist sport, but a sense of accomplishment in
terms of skill and physical endurance in conquest of nature red
in tooth and claw feature strongly.
43,44
There is still limited direct
research on why certain people engage in hunting for amuse-
ment, the ‘‘thrill of the chase,’’ or a ‘‘sense of achievement.’’
45
Overall, the ethical debates about recreational hunting in conser-
vation science literature have been mostly dominated, implicitly
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or explicitly, by a set of western-normative ethical perspectives
(e.g., deontology, utilitarianism/consequentialism).
24–26,46
Given
that perceptions, actions, and relationships with nature are influ-
enced by a diversity of worldviews, ethical commitments, values,
knowledge systems, and power relations, there are calls for a
more pluralist approach to ethical debates such as this.
47
These
approaches acknowledge alternative western positions such as
virtue ethics and ethics of care (e.g., Santiago-A
´vila and Lynn
48
),
as well as other non-Western ethical frameworks—for example,
Ubuntu, Buddhism, Hinduism, and buen vivir (e.g., Mkono
49
and
Gairola
50
).
HUNTING RESEARCH TOPICS
To assess the current state of knowledge on recreational hunt-
ing, and the degree to which the dominant research topics relate
to the conservation and livelihood implications of hunting, we
conducted a comprehensive key-word search in the Web of
Knowledge Core collection to identify peer-reviewed literature
related to recreational hunting. The search string comprised 25
key words associated with multiple forms of recreational hunting
(see Supplemental experimental procedures). We included items
published from 1953 to mid-September 2020, resulting in 3,882
references. We screened this initial list of articles for relevance
based on the content of titles and abstracts. We considered
only scientific articles explicitly addressing regulated recrea-
tional hunting, therefore discarding articles focusing solely on
commercial and/or subsistence hunting. We retained only those
articles dedicated to terrestrial ecosystems and mammal and
bird species. This screening resulted in 1,342 unique references
that we considered for further analysis (see Data S1). The num-
ber of articles published each year has been steadily increasing
over time at a rate similar to the literature focusing on general
conservation science (Figure 1).
Given the large number of studies, we used a natural-language
processing approach to identify the dominant topics discussed
in the collected articles. We used latent Dirichlet allocation,
51
which is a statistical model that identifies broad topics by ranking
key-word patterns across manuscripts. Specifically, topics are
identified by detecting key-word patterns within articles and
automatically clustering groups of key words that best charac-
terize a set of articles. Words used to discuss a particular topic
across all articles tend to occur together more frequently when
compared with the rest of the words, while a particular article
can discuss more than one topic.
52
The optimal number of topics
represents the fewest topics that maximize the information
covered as close to the original text as possible
53
(see Supple-
mental experimental procedures for detailed methods). Topics
were identified based on the title and abstracts of articles in
our database (when available; n = 1,171), and we list the resultant
seven topics in Table 1.
Articles associated with topic 1 address issues around the se-
lective harvesting of individuals, usually in the context of trophy
hunting. Selective harvesting can be sustainable,
54
but if poorly
managed it can have important ecological, evolutionary, and
population consequences. For example, the selective harvesting
of bighorn Ovis canadensis trophy rams in North America trig-
gered an evolutionary response toward reduced body weight
and horn sizes over time because the phenotypic traits targeted
by hunters are heritable.
55
Selective harvesting at unsustainable
rates can also modify natural sex ratios,
56
especially in sexually
dimorphic species when only one of the sexes possesses the
traits desired by most hunters, or change population structure
when individuals are selectively removed before reaching the
age of sexual maturity.
57
Figure 1. Number of studies on recreational hunting published per year
(A and B) The trendline in red in (A) shows that the number of studies published each year has been steadily increasing over time; (B) shows the percentageof
overall studies published in biodiversity and conservation that focus on recreational hunting. The search included items published from 1953 to mid-September
2020, hence the lower number of studies indicated in 2020.
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Hunting can alter the behavior of targeted species. Research
associated with topic 2 generally focuses on understanding
these behavioral responses. Research suggests that some spe-
cies, including the white-tailed deer Odocoileus virginianus,
58
red deer Cervus elaphus,
59
and American black bear Ursus
americanus,
60
might change their movement patterns in relation
to hunting pressure. Another frequent behavioral response of un-
gulates to hunting pressure is an increase in flight-initiation dis-
tances.
61
There is also evidence that hunting can cause species
to change their foraging behavior and habitat selection, and
induce physiological stress (e.g., Bryan et al.
62
). However, not
all species respond negatively to hunting (e.g., Neumann
et al.
63
), and research associated with this topic often focuses
on developing solutions to ensure sustainable hunting practices
by minimizing the impact of hunting on the target species’
behavior.
Articles associated with topic 3 explore the impacts of hunting
on population dynamics and provide assessments of drivers of
population status and trends, including hunting. Much of this
research focuses on estimating population parameters neces-
sary to understand temporal population dynamics, including
breeding success, age of first breeding, survival, and population
density (e.g., Devineau et al.
64
and Martinoli et al.
65
). Hunting
rates are also often reported (e.g., Angulo and Villafuerte
66
).
Research associated with this topic is essential to evaluate the
long-term sustainability of populations of hunted species and
to propose measures to ensure sustainability, such as the estab-
lishment of hunting controls and quotas (e.g., Dolman et al.
67
).
Topic 4 covers articles mainly dealing with hunting as a source
of, and population management tool for, invasive alien or conflict
species. Recreational hunting has been a driver for the introduc-
tion of game species—namely ungulates—in many regions of
the world; the red deer in New Zealand
68
and chital Axis axis in
Argentina
69
are two examples. These species can host zoonoses
and parasites,
70
and cause ecosystem impacts when their pop-
ulations expand in the absence of hunting. Articles associated
with this topic address these issues
71
and how hunting can
sometimes emerge as a tool to help control invasive or conflict
species’ populations and impacts.
72
Similarly, research in this
topic addresses how hunting can be a source of conflict with
wild species, namely predators, and a means used to control hu-
man-wildlife conflicts.
73
Research that addresses the health and ecotoxicological di-
mensions of hunting constitutes topic 5. Health concerns asso-
ciated with the consumption of wild meat have recently gained
additional attention in the context of the COVID-19 pandemic,
but similar concerns related to the consumption of game meat
have been a topic of research for many years. Examples include
analyzing the prevalence of pathogens in wild meat (e.g., Paul-
sen et al.
74
) or game meat compliance with health and safety reg-
ulations intended for global markets.
75
Lead pollution from hunt-
ing can also have detrimental consequences for the health of
wild animals and ecosystems. Much of the research associated
with this topic addresses contamination associated with the use
of lead ammunition in hunting, with potential consequences for
species, ecosystems, and even humans (e.g., Hampton et al.
76
).
Topic 6 includes research addressing hunters and their atti-
tudes, identities, and preferences, among other factors. Hunters
form a sizable and diverse group of wildlife-recreation enthusi-
asts, including people with different interests, skills, education,
views, and various degrees of specialization and engagement.
77
Much of the research associated with this topic describes these
characteristics, often in the context of the economic importance
of hunting. This includes understanding preferences for hunting
experiences (e.g., Schroeder et al.
78
), evaluating harvest expec-
tations (e.g., Bradshaw et al.
79
), exploring hunter identities and
views on different approaches to hunting,
80
or assessing willing-
ness to pay or travel for hunting.
81
The economic, social, cultural, political, and ethical dimen-
sions of hunting are the focus of research linked with topic 7.
Table 1. Seven dominant topics discussed in the recreational-hunting literature
Topic Description Top 10 associated key words
No. of articles as
dominant topic
1 Ecological, evolutionary, and population
consequences of selective harvesting
harvest, population, male, rate, age, size,
female, effect, mortality, survival
221
2 Ecological and behavioral consequences of
hunting, including activity and movement
patterns
area, human, habitat, density, study, forest,
pressure, effect, increase, site
158
3 Population dynamics in response to
hunting, including assessments of survival
rates, reproductive rates, and population
trends
management, population, model, estimate,
datum, control, base, approach, method,
potential
122
4 Hunting as a source of and management
tool for invasive alien or conflict species
deer, population, specie, wild, animal, red,
number, high, decline, region
150
5 Health and toxicological dimensions of
hunting
lead, bird, waterfowl, season, shoot, winter,
high, number, source, breed
157
6 Social dimensions of hunting, including
hunter attitudes, identities, and preferences
hunter, game, survey, recreational, wildlife,
bear, study, hunt, group, provide
159
7 Economic, social, cultural, political, and
ethical dimensions of hunting
conservation, trophy, wildlife, lion, local,
economic, tourism, community, benefit,
resource
204
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Much of this research highlights how hunting can provide a range
of contributions, both positive and negative, to humans and na-
ture. For example, it can be a source of revenue, which can be
used to support local economies
82
and the conservation and
restoration of threatened species and habitats.
14
Certain hunting
practices are also considered part of local cultural heritage in
different regions and therefore have deep cultural value.
83
On
the other hand, hunting can also be a source of conflict, owing,
for instance, to issues associated with land tenure and benefit
access by local people.
84
More broadly, hunting is also the focus
of heated political and ethical debates that permeate the peer-
reviewed literature. Articles debating the ethical and political di-
mensions of hunting, particularly trophy hunting (e.g., Hsaio
85
),
are included in this topic.
GEOGRAPHIC DISTRIBUTION OF HUNTING RESEARCH
To assess the geographic distribution of peer-reviewed knowl-
edge generation on recreational hunting, we considered where re-
searchers were affiliated, where their research took place, and
which topics were commonly researched inthese places. The au-
Figure 2. Geographic distribution of
recreational hunting research
(A and B) The maps show the country contribution
(as number of studies) to research, based on au-
thors’ affiliations of the studies (A) and countries
where the studies were done (B).
thors of the identified papers had affiliations
from 87 countries across all continents
(Figure 2A), while 147 countries across all
continents were researched in these
studies (Figure 2B). The United States (first),
Canada (second), and Spain (third) had
both the most articles that reported author
affiliations in these countries and the most
articles that had a research focus in these
countries. North America, Europe, and Af-
rica were the most researched continents,
although this might be partially an artifact
of our review only including studies pub-
lished in English. Research focusing on Af-
rica and Asia was done by authors based
in Africa or Asia, respectively, as well as
by authors based in Europe, North America,
and Oceania (Figure 3). Research focusing
on North America, Europe, Oceania, and
South America was instead mainly done
by authors based in those continents.
Geographic differences are evident
among the dominant research topics
(Figure 4 and Table 1). While all topics
were researched across Africa, North
America, and Europe, topic 7 (social
and ethical dimensions) was more com-
mon in Africa, while topic 6 (hunter per-
ceptions) was more common in North
America and topic 4 (population man-
agement) was more common in Europe. There were not
enough studies from the other continents to provide meaning-
ful continent-specific results.
TARGETED SPECIES
To identify the focal species in studies on recreational hunting,
we extracted species information (where present) from the arti-
cles (review papers were excluded to avoid repetition of studies).
There is a notable focus on certain species groups in the litera-
ture. Eight family groups were the focus of over 50 research ar-
ticles: Cervidae (n = 297 papers), Bovidae (169), Anatidae
(130), Felidae (117), Phasianidae (69), Suidae (68), Canidae
(64), and Ursidae (62).
Just 12 species were the focus of 50 or more studies each,
comprising 37% of all studies mentioning targeted species
(Figure 5A). There was a strong bias toward mammals (74% of
studies) compared with birds (26% of studies; Figure 5B). Eleven
of the 12 most commonly studied species were large mammals,
and just two are considered threatened (lion and leopard Pan-
thera pardus), with the remainder listed as Least Concern by
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the International Union for the Conservation of Nature (IUCN)
(Figure 5). These 12 species are either widely hunted in Europe
and North America or a target of trophy hunting in Africa. The
three most-studied species—red deer, white-tailed deer, and
wild boar Sus scrofa—have all been introduced for hunting in
several locations globally (e.g., wild boar and red deer intro-
duced to the Americas and Oceania; white-tailed deer intro-
duced to Europe and New Zealand), and hunting is now used
as a means of controlling their populations.
17
We identified studies that assessed the impact of recreational
hunting on the population abundance of targeted species as one
measurable dimension of species conservation (excluding re-
views, hunting to control problem species, and models predict-
ing future impacts of hunting). Methods of assessment included
long-term camera surveys, transect surveys, monitoring of ra-
dio-collared individuals, mark-recapture, and official records.
Just 35 species had more than one study each assessing the
impact of hunting on abundance (Figure 6). This highlights that
Figure 3. Global connections of recreational hunting research
Cords represent the connections between the countries where the authors were affiliated and where the research was done. Arrows point towards the country in
which the study was done. S. Am., South America; AR, Argentina; AUS, Australia; DK, Denmark; FI, Finland; MZ, Mozambique; NZ, New Zealand; PL, Poland; PT,
Portugal; RU, Russia; UK, United Kingdom; ZM, Zambia; ZW, Zimbabwe.
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despite the large body of literature assessing the impacts of rec-
reational hunting on targeted species (Table 1, topics 1 to 3), few
studies explicitly assess the consequences for abundance
trends (a commonly used indicator of conservation effective-
ness,
86
excluding cases where the objective is to cause
population declines of problem species). Recreational hunting
consistently caused abundance declines for one species (rein-
deer Rangifer tarandus). Findings were variable for a third of
studied species, with some but not all studies concluding that
hunting caused declines in abundance. This variability in conclu-
sions could reflect the diversity in studied locations (Figure 2B)
and time periods (Figure 1). For the remaining species, there
was no evidence of negative impacts of recreational hunting
on abundance. As a note of caution, the few studies and preva-
lence of inconclusive results for many species means that a lack
of evidence of negative impacts on abundance cannot be inter-
preted as evidence of sustainability.
Most studies (89%) assessing population impacts of hunting
focused on mammals, and over half of these were on large Afri-
can mammals (Figure 6). The most-studied species were two
large predators—lion and leopard—which are hunted for tro-
phies in 12 and 10 countries, respectively.
11
The mixed results
for these species (some studies showing negative impacts and
others not) demonstrate differences among and within countries
regarding the regulation of trophy hunting and the appropriate-
ness of hunting quotas (topic 3).
87–90
Sustainability challenges
include unscientific bases for setting quotas, excessive quotas
and offtakes, lack of restrictions on the age of individuals that
can be hunted, and other issues of poor governance (e.g., uncer-
tain property rights, corruption).
91–93
In some cases recreational
hunting is not the main threat to species, but can exacerbate the
larger impacts from poaching and habitat loss (e.g., trophy hunt-
ing can result in population declines if high poaching pressure is
not considered when setting quotas).
94,95
Three carnivore species in Europe/Asia (brown bear Ursus arc-
tos, gray wolf Canis lupus, fox) and one in North America (puma
Puma concolor) similarly showed mixed results, as did two North
American bird species (bobwhite quail Colinus virginianus, Can-
Figure 4. Stacked bar chart showing the
relative percentage of studies assigned to
broad recreational hunting research topics,
per continent
Total number of studies per continent is shown
above each stacked bar. Topics were identified by
using a latent Dirichlet allocation technique and are
further explained in Table 1.
ada goose Branta canadensis), with some
studies finding a negative impact of hunt-
ing on abundance and others not (Figure 6).
IMPLICATIONS OF HUNTING FOR
NATURE CONSERVATION
A holistic assessment of the effects of rec-
reational hunting on conservation requires
considering both the direct effects of hunt-
ing on the sustainability of the targeted species, as well as the in-
direct effects of this hunting on ecosystems more generally. Spe-
cies can be overharvested through recreational hunting,
resulting in population declines (Table 1, topic 3), or hunting
can have evolutionary and behavioral consequences for the
target species (topics 1 and 2). Hunting can disturb other spe-
cies, ecosystems, and ecological processes (topic 4), or cause
environmental pollution and poisoning of other species due to
discarded ammunition (topic 5). However, the socioeconomic
benefits generated by hunting can contribute to a species’ con-
servation and even reintroduction into areas of previous extirpa-
tion, and support habitat conservation and restoration (topic 7).
Recreational hunting is also a tool for controlling overabundant
and problem species, which can minimize their negative impacts
on other species (topic 4).
The interactions between recreational hunting and biodiversity
conservation are often context dependent and necessitate the
careful management of positive and negative conservation
trade-offs.
90
For example, several studies highlight the conser-
vation benefits and costs that arise from trophy hunting (topic
7). On the one hand, legislation provides private landowners
and communities in some southern African countries with user
rights over wildlife on their land, allowing them to generate reve-
nues through sustainable-use activities.
96
This legislation,
together with the growing ecotourism and trophy-hunting mar-
kets and low viability of livestock farming in arid areas, has re-
sulted in a transition from livestock ranching to wildlife ranching
over large tracts of land in recent decades, helping restore indig-
enous flora and fauna.
82,96,97
Approximately 1,394,000 km
2
of
land is dedicated for trophy hunting in sub-Saharan Africa,
exceeding the area encompassed by national parks there.
11,98
The white rhino Ceratotherium simum is an interesting example
of a species whose population recovery became tractable via
trophy hunting. The opportunity to purchase rhinos from their
last remaining population in Hluhluwe-iMfolozi Park, and offer
limited trophy hunting at high prices, incentivized South African
landowners to conserve and trade rhinos, resulting in their rein-
troduction across more than 16,000 km
2
.
99
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On the other hand, evidence shows that unsustainable hunting
of lions and leopards in some places has posed (or continues to
pose) serious risks to the conservation of these species
(Figure 6), calling for policy interventions to make this hunting
more sustainable.
20,88
In some contexts (e.g., leopard in South
Africa), such policy interventions have been implemented.
90
Predators are also sometimes killed because they prey on spe-
cies that are the target of trophy hunting, and the proliferation
of fences around wildlife ranches can be detrimental to wider
conservation efforts.
100
Furthermore, the hunting of captive-
bred lions on private land in South Africa has come under heavy
criticism.
11
Recreational hunting underpins the North American model of
wildlife conservation whereby hunting is proposed as an ethical
and conservation-motivated tool.
10
Since 1937, fees from water-
fowl hunters
101
have made it possible to conserve or restore
more than 50,000 km
2
of habitat in some of the most important
areas for duck and geese conservation in the United States
and Canada.
102
In addition, recreational hunting has now
become the primary product of the land for many landowners
in the United States.
103
In North America, hunting of abundant
herbivores and carnivores for management purposes is consid-
ered mostly sustainable,
13,104,105
which is supported by our find-
ings for moose and American black bear (Figure 6). However,
research quantifying the impacts of hunting on populations is
lacking for many species, and negative impacts have also
been documented for puma and Canada goose (Figure 6) (e.g.,
Packer et al.
106
).
Europe has a long history of using hunting as a tool to sup-
port biodiversity conservation.
107
In Denmark, for example,
hunting is an important motivation for landowners to retain
and enhance multifunctional landscapes through afforestation
of previous farmland and the establishment or restoration of
lakes and ponds.
108
However, habitat management to benefit
hunted species is not always good for other species.
109
Hunt-
ing has created the conditions for the introduction of invasive
speciessuchassikadeerCervus nippon and cotton-tail rabbit
Sylvilagus floridanus in Europe (topic 4), some of which have
damaged ecosystems.
22
Similar to findings for Africa, the dy-
namic behavioral and population ecology of large predators
can present challenges for achieving sustainable recreational
hunting in Europe (topics 1 to 3)
110
(e.g., gray wolf, brown
bear, fox—Figure 6).
111,112
The benefits of hunting game birds
for biodiversity conservation vary across Europe.
113
Most ev-
idence suggests that habitat management for game birds is
positive for other species in agricultural systems (topic 7),
113
and hunters in Europe also help scientists collect data that
can be used to monitor biodiversity trends.
114
By contrast,
conservation concerns include the illegal killing of protected
predators by those interested in the production of game spe-
cies, and the release of farm-reared birds into the wild to
augment populations.
113
Figure 5. Number of studies focusing on the recreational hunting of specific species
(A and B) The 12 most-studied species (A), and the percentage of studies dedicated to mammals and birds (B). IUCN Red List threat status: LC = Least Concern;
VU = Vulnerable. Silhouette images are from phylopic.org and in the public domain, except for Ursus americanus (creazilla.com; Creative Commons Attribution
4.0; by B. Comix) and Ursus arctos (pixabay.org; free for commercial use [Pixabay-License]).
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Figure 6. Impact of recreational hunting on the
population abundance of targeted species
Depicted is the proportion of studies that found incon-
clusive evidence, evidence of population declines, or
evidence of no population declines. Number of studies is
indicated in parentheses next to species name; only
species with >1 study are included. IUCN Red List threat
status: LC = Least Concern; VU = Vulnerable; NT = Near
Threatened. *Small antelope refers to steenbok Raphi-
cerus campestris, oribi Ourebia ourebi, grysbok Raphi-
cerus sharpei, duiker Cephalophus sp. or Sylvicapra
grimmia, and dik-dik Madoqua kirkii. Silhouette images
are from phylopic.org and in the public domain, except:
Kobus vardonii,Tragelaphus oryx,Alcelaphus busela-
phus,Kobus ellipsiprymnus,Phacochoerus africanus,
Redunca arundinum (phylopic.org; Creative Commons
Attribution 3.0; by J.A. Venter, H.H.T. Prins, D.A. Balfour,
R. Slotow, and M. Keesey); Damaliscus lunatus
(phylopic.org; Creative Commons Attribution-NonCom-
mercial-ShareAlike 3.0; by Lukasiniho); Vulpes vulpes
(phylopic.org; Creative Commons Attribution-NonCom-
mercial-ShareAlike 3.0; by A. Caravaggi); Tragelaphus
strepsiceros (WikiMedia.org; Creative Commons Attri-
bution-Share Alike 4.0 International; by user ‘‘Six Plus by
Libe
´’’); Ursus americanus (creazilla.com; Creative Com-
mons Attribution 4.0; by B. Comix); Ursus arctos
(pixabay.org; free for commercial use [Pixabay-
License]).
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Evidence on the implications of recreational hunting for nature
conservation in other continents is more limited, but at least for
Asia and South America this might arise from a dearth of En-
glish-language publications. In Asia, evidence suggests that rec-
reational hunting might affect population dynamics and sex ra-
tios of harvested populations in several countries (topics 1 and
3; Figure 6).
115
By depleting herbivores, recreational hunting
can also indirectly affect the conservation of threatened carni-
vores.
116
However, in the absence of carnivores in the
ecosystem, recreational hunting is also used to control popula-
tions of abundant herbivore species (e.g., Japan), therefore
reducing damage to natural habitats (topic 4).
117
In Oceania, rec-
reational hunting is mainly done to control overabundant spe-
cies, to decrease crop grazing and competition with livestock,
and to help conserve native habitat by controlling invasive spe-
cies (e.g., red deer).
68
However, some of these invasive species
were introduced for hunting, making hunting both the cause and
the solution (topic 4).
68
IMPLICATIONS OF HUNTING FOR LOCAL PEOPLE AND
ECONOMIES
Just two of the seven main topics in the literature focus primarily
on the human dimensions of hunting, with one (Table 1, topic 6)
focusing on the hunters themselves (particularly in North Amer-
ica) and the other (topic 7) focusing on the socioeconomic
outcomes of (particularly trophy) hunting for local people and
economies (particularly in Africa—Figure 4). Worldwide, recrea-
tional hunting has been an integral component of people’s diets
and cultural heritage, supporting livelihoods and contributing to
food security.
118
In addition, hunting has been important to fos-
ter local social-ecological knowledge, to develop meaningful re-
lationships with nature and sense of place,
118
to elicit pro-envi-
ronmental behavior, and to enable the evolution of formal and
informal institutions among hunters.
119
Hunting is used to miti-
gate human-wildlife conflict by controlling populations of
predators and large herbivores that pose a threat to human
life, livestock, and agriculture (e.g., Delibes-Mateos et al.
120
),
although it is not always effective (e.g., wild boar;
121
topic 4).
However, recreational hunting can also have negative implica-
tions for the health of local people by exposing them to the
spread of zoonotic diseases,
74,75
and to contamination by lead
ammunition
76
both from direct contact with wildlife and from
the consumption of game meat (topic 5).
Recreational hunting can generate revenue that accrues to
landowners, state conservation agencies, governments, local
communities, and/or the private sector, with the potential
(although not always realized) to provide livelihood opportunities
and funding for conservation (topic 7). Some studies have quan-
tified the hunting revenue accrued to private landowners and
state conservation agencies (and means of enhancing this reve-
nue; topic 6) in North America,
102,122
Europe,
123–125
sub-Sa-
haran Africa,
11,82,98
and Oceania.
126–128
The contributions that
recreational hunting makes to broader economies in these
regions have also been estimated.
122,129
Largely unique to
sub-Saharan Africa, there is also a body of literature assessing
the revenue accrued to local communities via hunting on com-
munity land or payments to communities residing within or adja-
cent to state hunting areas.
130–132
By contrast, few studies have
assessed the economic contributions of hunting in Asia or Cen-
tral and South America (but see Aryal et al.
115
and Baur et al.
133
).
A commonly mentioned threat to hunting revenue sources in
southern Africa is the growing pressure to ban trophy hunting
(done predominantly by foreign hunters).
11
In North America
and Europe, by contrast, the declining number of local recrea-
tional hunters in recent decades is noted as a concern for reve-
nue generation (e.g., in the United States
134
and Europe
17,135
). In
addition to fewer hunters potentially reducing incentives for
habitat and wildlife protection on private land, declines in
waterfowl hunting in the United States are predicted to result in
28–139 km
2
less land being restored each year due to lower rev-
enues from hunting stamps.
136
The North American model of
wildlife conservation has also been criticized, however, for
focusing its rhetoric on hunters and wildlife managers (topic 6)
and excluding non-consumptive users, policy-makers, and other
conservation practitioners.
137
The negative consequences of
declining hunter numbers in Europe for the management of prob-
lem species have also been discussed.
17,135
The contribution of recreational hunting to livelihoods and con-
servation is also influenced by the costs of conserving wildlife
(and who bears these costs; topic 7).
11
For example, trophy
hunting is estimated to generate US$138–1,091 km
2
across
sub-Saharan Africa, while estimates of the funding requirements
for effective management of protected areas are $460–
$2,048 km
2
.
138
Where wildlife authorities depend heavily on
hunting income to fund strict protected areas, the revenue can
be insufficient to manage hunting blocks or to provide meaning-
ful contributions to local communities.
132,139
The limited viability
of trophy hunting in some regions relates to the few remaining
trophy species, European Union bans on imports of trophies,
as well as increased costs of mitigating threats related to poach-
ing, agricultural encroachment, and growing climate insecu-
rity.
131,140
Elsewhere, while wildlife-based tourism (the primary
alternative wildlife land use) is limited to areas that combine
good infrastructure and abundant wildlife or spectacular scen-
ery,
141
trophy hunting can enable biodiversity conservation to
be a viable land use across many community conservancies
130
and private ranches.
142
There can be important power dynamics influencing who ben-
efits from hunting and how. Biodiversity in general, and hunters’
preferred species in particular, have been increasingly commod-
ified (i.e., neoliberal conservation), often through the imposition
of governance models that might not be suited to local con-
texts.
84,143
In some regions in sub-Saharan Africa, for example,
recreational hunting secured property rights and hunting quotas
while subsistence hunting by indigenous and local people was
banned and considered to be poaching, potentially leading to
social conflicts and compromising local support for conser-
vation.
84,144
Different initiatives have aimed to address these issues by pro-
moting the participation of local people in the governance of nat-
ural resources, including community-based natural resource
management projects in southern Africa.
145
Positive outcomes
include the creation of employment opportunities, increased
stakeholder collaboration, the emergence of local co-manage-
ment institutions, the engagement of communities in policy mak-
ing and monitoring, and improvement of local people’s attitudes
toward wildlife.
145,146
However, power dynamics can also be a
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challenge for these initiatives,
84,147
with imbalances in equitable
distribution of benefits, market challenges, and dependence on
tourism investors and trophy-hunting companies.
148
Failing to
address these issues in a fair way with communities can poten-
tially result in people’s disempowerment and active protests to
programs.
84
A challenge to understanding the implications of recreational
hunting for local people, and how this interacts with conserva-
tion, is the lack of clear counterfactuals—would local people
and nature have been better or worse off or the same in the
absence of recreational hunting? Historic and recent bans on
trophy hunting allow for one form of counterfactual analysis,
although such research is limited to a few examples (we there-
fore urge caution when attempting to extrapolate these findings
to other contexts). In Botswana, a hunting ban implemented in
2014 has been reported to affect local communities by reducing
income, employment opportunities, and access to game meat
(reducing food security).
145
This ban was subsequently lifted in
2019. In the case of Kenya where a ban on trophy hunting has
been in place since the late 1970s, there is potential support
for the return of trophy hunting to alleviate human-wildlife conflict
and enable access to benefits from hunting tourism.
149,150
Finally, assessments of the potential effects of hypothetical
bans on trophy hunting in Namibia and South Africa have re-
vealed unintended negative consequences for biodiversity con-
servation, economies, and livelihoods.
130,146,151
Overall, hunting practices and policies are highly context
dependent and are driven by a diversity of values, needs, identi-
ties, and worldviews. Despite this, existing research is limited in
quantifying local perceptions, experiences, power relationships,
and meanings on recreational hunting and its main stakeholders
at different scales.
147
A plural-valuation approach (e.g., Zafra-
Calvo et al.
152
) could contribute to recognizing and integrating
diverse value systems, needs, and ethical positions to foster
dialog, share understanding, and identify socially acceptable
hunting practices that would maximize the benefit to people
and nature.
OUTLOOK AND CONCLUSIONS
While the literature on recreational hunting is extensive, there are
clear biases toward certain continents, species, and topics, and
in many cases to species or populations that are not under immi-
nent threat of extinction (Figures 5 and 6). The focus on large
mammals reflects a common bias in conservation research.
153
This research bias also mirrors a focus on large charismatic
mammals in the literature on ethical concerns about recreational
hunting, and trophy hunting in particular (e.g., Vucetich et al.,
24
Batavia et al.,
25
and Ghasemi
46
). Surprisingly, despite the exten-
sive literature on recreational hunting, the evidence to address
some of the most pressing academic and societal questions
(e.g., when is recreational hunting sustainable, and who benefits
from it?) is still limited. Assessing the role of recreational hunting
in diverse social-ecological systems is critical to informing equi-
table ecosystem conservation and restoration approaches that
provide ecosystem services according to local people’s values
and needs.
While hunting is commonly mentioned as a threat to species
listed in the IUCN Red List,
2
it is not always clear whether the
threat is from legal forms of hunting or from poaching (i.e., illegal
hunting). Many studies research the impacts of recreational
hunting on the ecology and evolution of the targeted species.
Fewer assess the consequences for the abundance of species
of conservation concern, differentiating between the impacts
caused by recreational hunting versus poaching. Such assess-
ments are needed to quantify the implications of recreational
hunting for the sustainability of targeted species in diverse con-
texts. Similarly, it remains important to consider the interactions
between recreational hunting and other anthropogenic threats to
biodiversity (e.g., habitat loss and climate change), because syn-
ergistic processes between hunting and such threats could elicit
population declines even in cases where hunting is considered
sustainable. Such research should also focus on less
charismatic species that are covered poorly in the current
peer-reviewed literature, as well as assessing the impacts of rec-
reational hunting on non-targeted species and ecosystems (e.g.,
by better investigating the consequences of hunting large carni-
vores on predator-prey relationships and their cascading
ecosystem effects).
Beyond assessing the implications of recreational hunting for
the conservation of targeted species, our analysis reveals a
need for research that provides empirical evidence (ideally with
counterfactuals) for claims that hunting increases the quantity
and quality of ecosystem conservation. Many studies quantify
the revenues generated by hunting, but few demonstrate explicit
links between these revenues and conservation actions and out-
comes (e.g., funding to anti-poaching and reductions in poach-
ing, respectively). Similarly, research should focus on assessing
the contributions of recreational hunting areas to global and na-
tional biodiversity conservation objectives, and the quality of
management and its impact on biodiversity conservation in
these areas. This would provide a better proxy for assessing
the role of recreational hunting in supporting biodiversity conser-
vation, beyond the simple metric of total area that is conserved
on land dedicated to recreational hunting.
Another important topic of research we identified concerns the
health and toxicological dimensions of recreational hunting. The
ongoing COVID-19 crisis has renewed attention on zoonotic dis-
eases originating from wild animals. On the one hand, recrea-
tional hunting could increase the risk of human exposure to zoo-
notic diseases.
74,75
On the other hand, reduced tourist hunting in
the absence of revenue-generating alternatives could result in
local people relying more heavily on bushmeat consumption
and thus increasing the risk of future pandemics.
154
Effective
conservation of species and habitats has also been directly
linked to decreases in the number of viruses that animals share
with humans, meaning that recreational hunting could minimize
the risk of future pandemics in contexts where it contributes to
funding ecosystem conservation and restoration.
155
Future
research needs to investigate the potential connections between
recreational hunting and zoonotic disease risks, accounting for
these complexities.
While there is a large body of research focused on setting and
assessing hunting regulations such as quotas to promote sus-
tainability, there is comparably little research investigating the
role of local institutions (i.e., rules, norms, and traditions) in pro-
moting hunting that is both sustainable and beneficial to local
people (but see Bollig,
97
Tsas-Rolfes,
119
and Dowsley
156
). There
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is considerable potential for the common-pool resource litera-
ture to contribute theoretical insights and analytical tools to un-
derstand the interface of social and ecological dimensions of
wildlife governance,
157
yet the intersection of wildlife studies
and commons scholarship is not well studied.
158
Neither are
there many studies that consider conservation and social impli-
cations of hunting concurrently, despite their obvious connec-
tions. There is a need to engage the commons literature and
apply a social-ecological approach to understanding the con-
texts in which recreational hunting can provide benefits for
both people and nature.
Furthermore, there is a need to assess the moral complexity of
recreational hunting and to consider the perceptions, experi-
ences, power relationships, and meanings of recreational hunt-
ing for people in diverse contexts. Raising (self-)awareness of
these ethical issues among the scientific community and the
general public would contribute to the discussion, facilitating
dialog and shared understanding of the complexity and context
dependency around recreational hunting. Promoting legislation
in diverse sociocultural contexts in reaction to western-oriented
values without adequately consulting local people could have
detrimental effects on the hunted species and local people until
sustainable alternatives to recreational hunting are identified. It is
also important to determine whether benefits from hunting make
people more tolerant of the costs of living with biodiversity, and
thereby create an incentive for conservation among local com-
munities,
5
recognizing that benefits and costs are not only
monetary.
The decreasing numbers of hunters in Europe and the United
States, the increasing calls to ban trophy hunting in Africa, and
the COVID-19 pandemic all demonstrate the danger of conser-
vation and local livelihoods relying heavily on revenues from
tourism. Therefore, it is increasingly evident that alternative or
at least additional mechanisms for funding conservation need
to be identified.
154
Research could also assess whether recrea-
tional hunters would be willing to pay more to compensate for
diminished demand for hunting while also reducing pressure
on targeted species.
Our results should be considered with the following limitations
in mind. First, we only focused on peer-reviewed articles avail-
able in English, while many contributions on recreational hunting
are found in gray literature and in other languages. For this
reason, we consider our work as a first step toward a more
comprehensive review of all literature. Second, the breadth of
the literature and dominant topics prevented us from doing
meta-analyses to quantify the evidence around each topic. We
suggest that the topics we identified in this study could serve
as a basis to do more focused, topic-specific systematic litera-
ture reviews and other analyses. Future studies doing similar as-
sessments should now focus on the literature devoted to ‘‘sub-
sistence hunting’’ and ‘‘commercial hunting.’’ Future studies
should also attempt to define different types of hunting based
on motivations and regulations.
In conclusion, our assessment of the global peer-reviewed
literature on recreational hunting highlights that empirical evi-
dence on the contribution of recreational hunting toward meeting
both biodiversity and social objectives does not yet adequately
account for the diversity of contexts and worldviews. The need
for such evidence is urgent given declining numbers of recrea-
tional hunters in some regions and increasing opposition to tro-
phy hunting in others. The research agenda we propose to
address this knowledge gap emphasizes the importance of
considering the influence of local social-ecological dynamics
on hunting outcomes, and the voices of the people co-existing
with wildlife.
SUPPLEMENTAL INFORMATION
Supplemental information can be found online at https://doi.org/10.1016/j.
oneear.2021.01.014.
ACKNOWLEDGMENTS
E.D.M., G.C.-C., C.F., A. Hausmann, and R.K. thank the European Research
Council for funding under the European Union’s Horizon 2020 Research and
Innovation Program (grant agreement #802933). H.S.C. and A. Haukka thank
the Kone Foundation for personal grants. H.S.C. also holds a Jennifer Ward
Oppenheimer Research Grant. A. Haukka also thanks the University of Helsinki
for an Early Career Grant to E.D.M. R.A.C. thanks the University of Helsinki for a
personal grant to E.D.M. All authors would like to thank M. Festa-Bianchet and
two anonymous reviewers for comments that helped us improve our manu-
script.
AUTHOR CONTRIBUTIONS
Original Idea, E.D.M. and C.J.A.B.; Formal Analysis and Investigation Soft-
ware, R.A.C. and R.K.; Data Curation, E.D.M., H.S.C., R.A.C., and A. Haukka;
Visualization, E.D.M., H.S.C., R.K., C.F., and A. Haukka; Writing – Original
Draft, E.D.M., H.S.C., R.A.C., G.C.-C., and A. Hausmann; Writing – Review
and Editing, E.D.M., H.S.C., R.A.C., G.C.-C., C.F., A. Haukka, A. Hausmann,
R.K., and C.J.A.B.
DECLARATION OF INTERESTS
The authors declare no competing interests.
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