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What does trophy hunting (selective hunting for recreation) contribute to wild lion conservation? Macdonald (Report on Lion Conservation with Particular Respect to the Issue of Trophy Hunting. WildCRU, Oxford, UK, 2016) summarises what we know. We identify unknowns, gaps in the knowledge that inhibit conservation planning, including: the causes of lion mortality, the amount of land used for lion trophy hunting, the extent to which trophy hunting depends on lions for financial viability, and the vulnerability of areas used for hunting to conversion to land not used for wildlife, if trophy hunting ceased. The cost of reversing biodiversity loss exceeds income from tourism, including hunting. New financial models are needed, particularly in view of the expanding human population in Africa.
Lions Panthera leo, iconic symbols of the African wilder-
ness, are in trouble. The latest International Union for
Conservation of Nature (IUCN) estimates suggest a popu-
lation of 23000–39000 (probably closer to the lower es-
timate), a decline of at least 43% between 1993 and 2014,
approximately three lion generations (Bauer et al. 2015).
Lions have been extirpated from at least 92% of their
historic range. According to the 2016 IUCN Red List
Assessment, in Africa, lions are almost certainly extinct
in 15 countries, possibly extinct in another seven, and
now occur in only 25 countries (Bauer et al. 2016). Lion
decline may be even more severe than estimated by the
IUCN (Anonymous 2006a, b) based on well- known popu-
lations which tend to be in a better state. Only six popu-
lations have more than 1000 lions (Selous- Niassa,
Serengeti- Mara, Kavango- Zambezi, Greater Limpopo,
Lions, trophy hunting and beyond: knowledge gaps and why
they matter
David W. MACDONALD* Wildlife Conservation Research Unit, Department of Zoology, The Recanati-
Kaplan Centre, University of Oxford, Tubney House, Tubney, Oxon OX13 5QL, UK.
Andrew J. LOVERIDGE Wildlife Conservation Research Unit, Department of Zoology, The Recanati-
Kaplan Centre, University of Oxford, Tubney House, Tubney, Oxon OX13 5QL, UK.
Amy DICKMAN Wildlife Conservation Research Unit, Department of Zoology, The Recanati-Kaplan
Centre, University of Oxford, Tubney House, Tubney, Oxon OX13 5QL, UK.
Paul J. JOHNSON Wildlife Conservation Research Unit, Department of Zoology, The Recanati-Kaplan
Centre, University of Oxford, Tubney House, Tubney, Oxon OX13 5QL, UK.
Kim S. JACOBSEN Wildlife Conservation Research Unit, Department of Zoology, The Recanati-Kaplan
Centre, University of Oxford, Tubney House, Tubney, Oxon OX13 5QL, UK.
Byron DU PREEZ Wildlife Conservation Research Unit, Department of Zoology, The Recanati-Kaplan
Centre, University of Oxford, Tubney House, Tubney, Oxon OX13 5QL, UK.
Mammal Review (2017) © 2017 The Mammal Society and John Wiley & Sons Ltd
Africa, economics, ethics, lions, trophy hunting
*Correspondence author.
Submitted: 3 April 2017
Returned for revision: 15 May 2017
Revision accepted: 12 June 2017
Editor: DR
doi: 10.1111/mam.12096
What does trophy hunting (selective hunting for recreation) contribute to wild
lion conservation? Macdonald (Report on Lion Conservation with Particular
Respect to the Issue of Trophy Hunting. WildCRU, Oxford, UK, 2016) sum-
marises what we know. We identify unknowns, gaps in the knowledge that
inhibit conservation planning, including: the causes of lion mortality, the amount
of land used for lion trophy hunting, the extent to which trophy hunting de-
pends on lions for financial viability, and the vulnerability of areas used for
hunting to conversion to land not used for wildlife, if trophy hunting ceased.
The cost of reversing biodiversity loss exceeds income from tourism, including
hunting. New financial models are needed, particularly in view of the expanding
human population in Africa.
Mammal Review ISSN 0305-1838
2Mammal Review (2017) © 2017 The Mammal Society and John Wiley & Sons Ltd
D. W. Macdonald et al.Unknowns in lion trophy hunting
Katavi- Ruaha and Kgalagadi). About half of the remaining
wild populations comprise fewer than 100 individuals
(Dickman et al. submitted). Lion populations are in crisis,
due primarily to the loss and degradation of habitat, loss
of prey, and conflict with people (Anonymous 2016a),
pressures exacerbated when they are small, isolated, and
poorly managed (Anonymous 2006a, b, Henschel et al.
Public interest in lion conservation was stimulated in
July 2015 with the killing of a well- known lion nicknamed
‘Cecil’ (Macdonald et al. 2016a). Fierce debate has since
raged over whether trophy hunting is good or bad for
lion conservation. To be clear, trophy hunting ‘generally
involves the payment of a fee by a foreign or local hunter
for a hunting experience, usually guided, for one or more
individuals of a particular species with specific desired
characteristics (such as large size or antlers). The trophy
is usually retained by the hunter and taken home’
(Anonymous 2016a).
This debate would be informed by identifying the con-
ditions where trophy hunting contributes to lion conserva-
tion, if indeed such conditions prevail anywhere. Whether
trophy hunting lions is ethically acceptable is a distinct
debate, which we enlarge on below. Some hold trophy
hunting in such moral repugnance that any benefit to
conservation is insufficient to justify it. This view may
come to prevail, and perhaps a majority of the Western
public already holds it (although the balance of opinion
probably varies from place to place, notably between the
West and lion range countries). Until the part played by
trophy hunting is known and, as necessary, alternative
means of financing lion conservation are in place, we
have defended a more utilitarian population- based per-
spective, arguing that the cost of implementing such a
moral imperative may be too high if no better alternative
for lion conservation is available (Macdonald et al. 2016b).
We support the cessation of trophy hunting where it is
clearly inimical to conservation, and its reform where it
is better for conservation than any viable alternative land
use. While lion hunting exists, it should, at least, be sus-
tainable. An account of the evidence base for assessing
its sustainability is presented by Macdonald (2016). In
compiling that evidence base, we were, however, thwarted
by a surprising lack of information on several important
issues. In the much, but perhaps unfairly, mocked words
of the former US politician Donald Rumsfeld, there are
knowns, known unknowns, and unknown unknowns.
We identified a conspicuous knowledge gap concerning
the causes of lion mortality. At both national and regional
scales, trophy hunting of wild lions has been ranked rela-
tively low as a threat (Anonymous 2006a, b; we do not
consider ‘canned’ hunting, in which captive- bred lions are
hunted, as it has little relevance to wild lion conservation).
However, there is evidence that over- hunting has reduced
lion numbers at national scales – such an effect was par-
ticularly clear in Tanzania (Packer et al. 2011). In unfenced
reserves, large- scale population growth rates are lower in
the presence of trophy hunting (Packer et al. 2013). It is
also clear that badly regulated hunting can be locally
damaging (Caro et al. 2009, Packer 2015, Creel et al.
Trophy hunting is not the only reason lions are killed.
How the number of lions killed by trophy hunting com-
pares with those killed by snaring or human- lion conflict
is known only for a few localities (e.g. Loveridge et al.
(2016), and conventional ecological methods for estimating
mortality have been found to underestimate rates of illegal
killings, such as poaching, relative to legal killings, for
example trophy hunting (Treves et al. 2017). Mortality
due to conflict with local people may be orders of mag-
nitude greater than that due to international trophy hunt-
ers: in Tanzania’s Ruaha landscape, at least 37 lions were
killed in 2011 due to conflict, in an area of less than
500 km2, making the offtake over 100 times higher than
the recommended maximum offtake for a trophy hunting
area (Dickman in prep.). Recently, concern over lions be-
ing poisoned as an incidental outcome of attempts to
disguise elephant poaching has gained prominence (Sandhu
2016). There are other places (e.g. Hwange; Loveridge
et al. 2016) where trophy hunting is the main cause of
mortality for adult male lions. However, in many places
the balance of these factors remains unknown, although
it is suspected that throughout Africa many more lions
die due to conflict than are killed by trophy hunters
(Anonymous 2016a).
Where trophy hunting occurs, its mortality is probably
additive. It can also lead to a cascade of indirect mortality
through social perturbation (the perturbation effects of
other sources of mortality have not been studied). But
even where other sources of mortality predominate, it is
theoretically possible for a small amount of additive mor-
tality to tip the balance from a scenario where a popula-
tion is stable or increasing to one where the population
growth rate is negative. Creel et al. (2016) concluded that
for trophy hunting to be sustainable under the conditions
that most lions experience, total mortality needs to be
reduced. Where other sources of mortality dominate, tack-
ling them is likely to be the priority. It is clear that there
will be many places where focussing on a single threat
to lions, whether trophy hunting or any other cause, will
be inadequate for effective conservation. A holistic ap-
proach, considering all the threats and their interconnec-
tions, is most likely to succeed.
As with photo- tourism, trophy hunting can protect
wildlife by providing an economic reason for land being
maintained under a wildlife- based land use. Income
Mammal Review (2017) © 2017 The Mammal Society and John Wiley & Sons Ltd
Unknowns in lion trophy huntingD. W. Macdonald et al.
from trophy hunting can be significant for government
bodies responsible for managing wildlife – the Wildlife
Division in Tanzania, for example makes 60% of its
income from trophy hunting licence fees (Estes 2015).
Major unknowns therefore include the likely, but in-
conclusively demonstrated, positive roles of trophy hunt-
ing in creating economic incentives for wildlife- based
land use, whether on private, state or communal land.
It may also provide an anti- poaching presence in wildlife
areas, together with general management such as main-
tenance of boreholes. How widely land managed for
hunting is well managed for non- hunted species is also
poorly understood.
The continued availability of land for lions is clearly
crucial for their conservation. Our review of the evidence
led to the conclusion that trophy hunting’s greatest con-
tribution to lion (and wider) conservation lay in providing
an incentive to retain wildlife habitat that might otherwise
be lost to agriculture or pastoralism (Macdonald 2016).
It is vital, therefore, to know how much lion habitat there
is, where it is, how many lions it supports and how it
is managed. Large areas of wilderness are used for lion
hunting, but there are few recent estimates of precisely
how much. Lindsey et al. (2013) estimated that lions were
hunted in around 558000 km2, which comprised 27–32%
of the lion range in countries where they were hunted.
There have been no published updates since Botswana
banned lion hunting in 2008 (and all hunting in public
areas in 2014), or since Zambia imposed a lion hunting
moratorium between 2013 and 2016. Also, since Lindsey
et al.’s estimate, human encroachment has caused losses
of hunting land: Packer (2015) estimates that 40% of
hunting blocks in Tanzania have been abandoned in the
last decade. Hunting blocks elsewhere have also been
abandoned after becoming depleted and unviable (Lindsey
et al. 2016). If not used for hunting, a lot of that land
would be likely to be lost to wildlife, by for example
being converted to agriculture or livestock grazing. In some
areas, where economic forces did not prompt conversion,
lions and their prey might recover when hunting stopped,
leading to restoration of hunting at some point.
We make two interim conclusions: first, trophy hunting
should be strictly regulated to ensure that it does con-
tribute to lion conservation, including by the maintenance
of habitat (Macdonald 2016 makes clear how this can be
achieved). Second, where lion hunting is disallowed by
national law or rendered financially unviable (by import
bans for example), alternatives must be found to ensure
that its contribution to habitat preservation is replaced
– this is the difficult bit, and the one worst bedeviled by
unknowns. The substituted institutions will need to effect
more than habitat protection, by preventing poaching, for
example. It is crucial to distinguish between scenarios
where trophy hunting of lions alone is stopped and those
where there is a general cessation of trophy hunting. Many
hunting areas may not be financially dependent on lions,
but a further unknown is whether a cessation of lion
hunting would be followed by extended restrictions on
the hunting of other charismatic and threatened species.
Lindsey et al. (2017) demonstrate that, with or without
hunting, many areas have insufficient funds for effective
An understanding of the mechanisms whereby trophy
hunting affects lion populations requires monitoring, and
knowledge not just of population size, but also of the
density of individuals eligible for hunting. Appropriate
methodologies are available (e.g. Funston et al. 2010,
Broekhuis & Gopalaswamy 2016, Elliot & Gopalaswamy
2016). Macdonald (2016) shows how an adaptive manage-
ment system can ensure that departures from sustainable
offtakes can be rectified. Thus, while there is scope for
refining methods of counting carnivores (Gopalaswamy
et al. 2015), useful methods exist – the dangerous unknown
is ignorance of the numbers of lions, largely due to the
practicalities of who is going to pay for such
In principle, calculating the mortality lion populations
can withstand, from trophy hunting or any other source,
is straightforward. There are area- based and density- based
harvesting models of sustainable offtake. These could be
refined to account for intra- specific variation in lion den-
sity, and other threats. For example the figure recom-
mended by Packer et al. (2011) for offtake of 0.5 lions
per 1000 km2, while intended to be precautionary, does
not account for variation in lion population density.
Furthermore, it would be useful to quantify the interac-
tions between mortality factors (e.g. trophy hunting, snar-
ing, and conflict) some of which (e.g. snaring) are
non- specific. Perturbation effects (Tuyttens & Macdonald
2000) on lion demography resulting from trophy hunting
are well documented (Loveridge et al. 2007), and there
is evidence that such effects exacerbate human- lion conflict
(Loveridge et al. in prep).
It is difficult to predict what would happen to hunted
lion populations if hunting was stopped. Would photo-
tourism substitute, or some other regime that was not
wildlife- friendly? Hunting may, in general, be less beneficial
4Mammal Review (2017) © 2017 The Mammal Society and John Wiley & Sons Ltd
D. W. Macdonald et al.Unknowns in lion trophy hunting
than photo- tourism for lion populations, but may none-
theless be better than nothing (Lindsey et al. 2017). There
are crucial unknowns from the viewpoints of both land
managers and governments. These include the extent to
which photo- tourism (itself vulnerable to insecurity and
global economic forces), or other non- consumptive uses
can substitute for trophy hunting, and crucially, how lion
populations would fare following conversion. How much
current trophy hunting land is suitable for conversion to
photo- tourism is uncertain. Lindsey et al. (2006) argue
that certainly not all of it is, and question whether there
is sufficient demand to supply visitors to these areas.
Under what conditions is trophy hunting land converted
to less wildlife- friendly uses such as agriculture, settlement,
mining, or pastoralism? The regulation and enforcement
of land ownership and land- use zoning are likely to be
influential. The key unknown here is how important lions
are for the profitability of trophy hunting operations rela-
tive to their value under different land uses. It is also
important to whom the different values of lions accrue
under different land- use systems. Progress is hampered
by inadequate political, legal, and governance instruments,
such that local people have no incentive to value wildlife
(Muposhi et al. 2016).
Amongst the most serious unknowns, then, are the
extents to which trophy hunting (and photo- tourism) does
or could provide sufficient financial incentive to retain
land under wildlife- based uses and under alternative uses,
and what factors influence this (and how the answers are
likely to change in the rapidly changing socio- economic
landscape of Africa and beyond). From the viewpoint of
governments, the associated expenditure and value- added
estimates of the economic contribution of trophy hunting
and the alternative land- uses should be compared, not
just the economic revenue directly attributable to each
activity, which is most commonly the practice in both
the academic literature and advocacy documents. Questions
of land- use transitions will also be affected by unknowns
such as the likely sequence in which measures against
lion hunting would be extended to other species, most
obviously elephants Loxodonta africana to whom, de facto,
it has already extended through the ban of imports in
important consumer countries, whereas the process of up-
listing leopards Panthera pardus already began under the
Endangered Species Act of the USA (Anonymous 2016a,
In 2012, before the restrictions on elephant hunting
and reduced lion quotas, Lindsey et al. (2012) made a
tentative prediction that a lion hunting ban would make
trophy hunting financially unviable in substantial areas of
the lion’s geographic range, with potential wider negative
impacts. Banning the hunting of species like leopards would
be likely to reduce viability in a wider area. Those authors
did not account for the cost of conservation in their per-
spective on sustainability. Questions about the proportion
of park and wildlife management budgets provided by
trophy hunting operators are generally unanswered (and
might usefully be posed of photo- tourist operators too).
Most National Parks in Africa would not be financially
viable without support from government, which often
comes at least in part from hunting revenues. It would
also be helpful to know how important trophy hunting
is for the financial viability of wildlife authorities.
Considering that in most African countries conservation
is underfunded (Lindsey et al. 2017), it would be useful
to know more about the comparative economics of trophy
hunting and photo- tourism.
A linked unknown is how lions would be tolerated if
they could not be hunted, but land was managed for
other wildlife uses, such as trophy hunting of their prey.
Lions could then impose a substantial cost – it has been
said that ‘game farming is incompatible with predators’
(Schneider 1990). The fate of lion populations, even under
non- consumptive land uses, is also uncertain, although
insights may soon be gleaned from Botswana where hunt-
ing was banned in 2015 (Macdonald 2016); there may
also be lessons, although certainly not simple cause and
effect, to be gleaned from Kenya, where trophy hunting
was banned in 1977 and where wildlife numbers declined
on average by 68% between 1977 and 2016, alongside
increases in human and livestock numbers that further
confound simple interpretations (Macdonald 2016, Ogutu
et al. 2016). Banning trophy hunting does not necessarily
lead to less killing of lions: Richard Leakey observes that:
Carnivores are being decimated… hunting has never been
stopped in Kenya, and there is more hunting in Kenya
today than at any time since independence. (Thousands)
of animals are being killed annually with no control…’
(Martin 2015).
Of 38 lion populations in non- hunting areas examined
for the latest IUCN Red List assessment, 58% were de-
clining, whereas of the seven populations examined in
hunting areas, potentially self- selecting and mainly fenced,
one (14%) was declining (Bauer et al. 2015). Comparing
trends is not straightforward. For example, hunted popula-
tions are likely to be depleted already, whereas well-
protected populations are generally closer to carrying
capacity and are therefore more likely to decline if pro-
tection wavers.
It is clear that evaluating how trophy hunting contributes
to lion conservation is compromised by lack of data. Here,
we turn to the current state of lion trophy hunting.
Mammal Review (2017) © 2017 The Mammal Society and John Wiley & Sons Ltd
Unknowns in lion trophy huntingD. W. Macdonald et al.
Macdonald (2016) sets out criteria under which lion hunt-
ing could be deemed sustainable. How many hunted
populations meet these criteria is unknown, as are the
conditions where trophy hunting is a conservation tool.
Although we know which populations are currently hunted,
we do not know how many of these depend on lion
hunting for their viability (the only estimate, from 2012
and before heavy quota reductions in Tanzania and several
other countries, is about 11%; Lindsey et al. 2012). Indeed,
for many management units, how many lions are hunted
annually is unknown. Monitoring of both populations and
hunting offtake is often poor; Macdonald (2016) concludes
that under these circumstances, precaution demands the
use of conservative age- based and area- based criteria when
allocating quotas.
Amongst the knowledge gaps that impair a compre-
hensive analysis of lion trophy hunting is the inadequacy
of statistics on exports. Improving the collection and or-
ganisation of data by the Convention on International
Trade in Endangered Species of Wild Fauna and Flora
was amongst several recommendations made by Macdonald
(2016). Others included a move to open auctions for
concession leases and to longer leases to discourage short-
termist mining of the natural resources (Brink et al. 2016).
Thus far we have focused on empirical data. However,
the evidence of these disciplines will be judged within a
wider set of societal ethics. There is no consensus, even
among conservationists, that a utilitarian perspective on
trophy hunting is the right one. Even the concept of
‘sustainability’, used above as a criterion of good manage-
ment, would be viewed by some as ethically questionable
when applied to lion hunting (or to any killing of animals
for ‘sport’). Also, we are mindful that while emotional
responses affect moral judgements (Nelson et al. 2016),
policies based principally on emotion could have perverse
consequences. Where an intention to improve lion con-
servation worsens it, perhaps even those implacably op-
posed to lion hunting on ethical grounds might favour
a ‘journey’ towards its cessation rather than a ‘jump’. As
Macdonald (2016) concluded ‘if society judged trophy
hunting lions unacceptable, but also concluded that it
benefited lion conservation, then this dilemma might be
approached via a journey to find ways of replacing the
benefits of hunting before jumping to end them’.
The day may not be far off, if it is not here already,
when much of society (at least outside lion range coun-
tries) regards lion hunting as being as unacceptable as,
for example bear baiting or child labour (Macdonald et al.
2016a). However, views widely held in the wealthy West
are often at odds with views within lion range countries,
where lions often impose severe costs (including man-
eating) on the people who live alongside them. Who has
the right to make decisions about trophy hunting? How
should the weight of opinions held on lion hunting in
countries without lions, such as the USA (which has a
thriving domestic hunting market), be ranked against the
opinions held in African countries where lions occur (and
where the financial consequences of a cessation of trophy
hunting might bite hardest)? These are all difficult issues.
It is clear, though, that if lion hunters aspire to be toler-
ated, they must demonstrate radical reform (and that may
not be enough). It is also clear that those who seek to
eliminate trophy hunting have either to acknowledge that
the possible subsequent loss of lions is a cost they are
prepared to pay, or to demonstrate an economically valid
alternative wildlife- based land use.
Trophy hunting, like almost everything else affecting con-
servation, is a moving target (and moving, like all aspects
of African conservation, heavily at the mercy of external
factors). Having reviewed what we know about lion trophy
hunting (Macdonald 2016), we thought it helpful to high-
light the unknowns here and why they matter. Our un-
derstanding of trophy hunting’s potential global significance
for the species is compromised by not knowing over how
much of the species’ range it occurs. Where trophy hunt-
ing does occur, the implications for lion conservation of
any change to the current system vary from place to place.
Where lion trophy hunting is run with sustainable quotas,
and where no viable wildlife- friendly alternative exists, its
removal seems likely to be negative for lion conservation.
But there are extensive areas where the implications of
the removal of trophy hunting for lion conservation are
uncertain, because we do not know the answer to ques-
tions like how much the industry’s viability depends on
lions, or if lions could persist after an alternative land
use was substituted.
Unknown threats to lions will surely change. The next
clutch will be spawned by changing societal, global eco-
nomic, demographic, and environmental factors. Trophy
hunting, and the prudence of relying on tourism to sup-
port conservation in Africa, might be considered minor
issues compared to the others jeopardising biodiversity.
The money needed to reverse biodiversity loss dwarfs that
likely to flow from any variant of tourism, including hunt-
ing, so new financial models to encourage coexistence with
nature must be found. Dickman et al. (2011) speculate
that mechanisms for converting global value to local ben-
efits provide one promising option. Beyond that, we cannot
predict how emerging markets and economies such as
Russia and China will influence the status quo. The most
6Mammal Review (2017) © 2017 The Mammal Society and John Wiley & Sons Ltd
D. W. Macdonald et al.Unknowns in lion trophy hunting
perilous unknowns of all include the consequences of an
estimated tripling by the year 2100 of the human popula-
tion of Africa. Whatever plan is put in place for the
conservation of lions and the rest of Africa’s wildlife, it
must accommodate the reality of nature living alongside
two billion people.
We thank H. Bauer, J. Vucetich, and D. Burnham for
comments, and gratefully acknowledge discussion with G.
Chapron, P. Lindsey, C. Packer, together with T. Hodgetts
and K. Somerville, and all those who commented on our
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... This research study was guided by ethical theory into four aspects of deontology, utilitarianism, and virtue ethics (Darimont et al., 2017;Benjamin 2020), which are widely used to explain sport hunting practice in wildlife conservation areas. The theory conceptualizes that sport hunting must not jeopardize wildlife populations, alter natural selection and ecosystem functioning, or diminish native biodiversity (Macdonald et al., 2017). ...
... According to Macdonald et al. (2017), sport hunting and wildlife conservation have both been part of human culture from the earliest times. As the World Bank Group (2019) notes, sport hunting does not always target problem animals. ...
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How sport hunting influences wildlife conservation in Uganda within the Lake Mburo Landscape.
... There is a widespread agreement that the impact of recreational hunting on wildlife depends on the socio-ecological context [1][2][3][4]. On the one hand, studies show that recreational hunting can positively affect wildlife and biodiversity conservation, e.g., [5][6][7][8][9][10][11][12]. ...
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Recreational hunting can have positive consequences on biodiversity conservation, but in many countries the number of hunters is declining. This downward trend threatens the sustainability of an important human activity that can be used as a tool for wildlife management and biodiversity conservation. On the other hand, in developed countries there is an upward trend in the number of female hunters. In this study, we analyzed women and men's hunting preferences in Spain, focusing our attention on the game species they were interested in. We found that female hunters were more interested than male hunters in hunting big game species. We discussed potential consequences of our results on the sustainability of hunting and biodiversity conservation.
Within the realm of social carnivores, lions (Panthera leo) are the sole representative of the large felids. While other Felidae may tolerate temporary associations with dependent offspring or receptive members of the opposite sex, lions are the only big cat where both males and females live in permanent social units. Each sex forms independent groups that come together to engage in a wide variety of social behaviors, including cooperative hunting, mutual defense of kills, cooperative territorial defense, and communal raising of young. Unique patterns of egalitarianism and flexible fission-fusion dynamics make this social structure distinctive among mammals. The bulk of our knowledge on the evolutionary drivers of lion sociality has been derived from extensive studies of populations inhabiting mesic and productive eastern African savanna systems. However, lions across Africa occupy a wide variety of habitat types, including arid deserts, seasonally flooded wetlands, and densely thicketed scrub. Comparison of lion behavior and interactions across their range reveals that their social strategies are highly plastic and adapt to maximize survival and fitness under prevailing local conditions. In this chapter, we first review the ultimate drivers of lion gregariousness based on research from eastern Africa and discuss how lions optimize individual fitness under constraints imposed by group-living. We then explore how variation in proximate drivers of social living (i.e., resource availability, intensity of inter- and intra-specific competition, and habitat structure) can shape the expression of social behavior. We end with a discussion of the social adaptations lions make to survive in increasingly human-dominated landscapes.
The most basic moral dilemma in sport hunting is the dispute between deontologists, arguing that animals have inalienable rights to life, and consequentialists, arguing that hunting can lead to less total suffering and the conservation of species and habitats. This dilemma has already been presented in the historical chapter, mainly in Chapters 2.9 and 2.10.What we will attempt to demonstrate in Chapter 8 is that deontology vs. consequentialism is not the only important conflict between paradigms of normative ethics in the trophy hunting discourse. What seems to be unique about the conflict over trophy hunting compared to the conflict over sport hunting is that there is less emphasis on the death of animals per se and more emphasis on the persons who cause said deaths. The emphasis is on the character, motivations, behaviour, and attributes of the hunter.Our claim, following from this observation, is that in order to understand the moral conflicts underlying the trophy hunting discourse, it is no longer enough to understand the obvious incompatibilities between deontology and consequentialism. We must also be open to the possibility of incompatibilities between virtue ethics (i.e. the character of hunters) and consequentialism.To illustrate this, we present in Chapter 8.1 and 8.2 our observations from Twitter, where comments to trophy photos were categorized as pertaining to either the character of hunters or animals/the death of animals. A far greater number of comments about the character of hunters were found, supporting the notion that virtue ethics play an important role in antihunting sentiments.
“Sport Hunting” is highly contentious and confusing, because it can have two meanings. One meaning of sport hunting is to hunt in a sporting way and give the animal a sporting chance. This is equivalent to the ideal of fair chase. The other meaning is to hunt for sport. It portrays hunting as competition and fun.In Chapter 6, we summarize the history of these terms – also discussed in Chapter 2 already – and discuss the meaning of sport hunting. We then discuss how fair chase is sought achieved by decreasing the power gap between hunter and prey through technological handicaps (like using a bow instead of a rifle – Chapter 6.1) and behavioural handicaps (like not shooting a deer on ice or in water or deep snow – Chapter 6.2).We then cover in Chapter 6.3 what we suggest could be an inverse relationship between fair chase and animal welfare. Bowhunting is one such example where, by making the hunt more difficult and therefore “fair,” evidence suggests that wounding rates and thereby animal suffering may increase.Finally, canned hunting is the practice of hunting animals that are fenced in, thus limiting their chance of escape. We discuss the very controversial issue of canned hunting in Chapter 6.4, as it is generally held up as an example of the opposite of fair chase. Plenty of hunting ranches in America, notably Texas, offer canned hunting, and captive-bred lions in South Africa for lion-petting tourism and canned hunting has until now been a big industry but seems to be shutting down. We cover all of this in 6.4.
This chapter investigates conservation claims and issues as they pertain to hunting. After a description of the major regulations governing trophy hunting imports and exports (5.1 Trophy Hunting Regulations), this chapter examines the conservation situation in two geographically different sections.The first section (5.2 Hunting and Conservation in Africa) concerns the stereotypical perception of trophy hunting. The trophy hunting situation and conservation issues are completely different in Africa than in Europe and USA and the problems are more diverse and complex. African hunting is more controversial because the species involved (lions, elephants, giraffes, etc.) are iconic, highly anthropomorphized, and sometimes endangered. Both species extinction and species overpopulation are issues in Africa. Corruption, poverty, poaching, and the West imposing their wildlife ideals on Africa are also important factors that we cover here.The second section (5.3 Hunting and Conservation in Eurasia and the Americas) concerns trophy hunting mostly in Europe and USA, where they have few natural predators of deer, so hunting is broadly considered the most feasible way of keeping deer populations from outgrowing carrying capacities of habitats. There are issues, however, with keeping populations down, because hunters favour bucks (because of trophy-fixations and ingrained fair chase ideals), and shooting bucks does little to keep populations in check. Also, shooting the wrong bucks or shooting them too early leads to genetic problems in the populations. These are the primary conservation issues that we discuss in a European- and US context.Other conservation-related matters discussed in this chapter are hunting and genetics, management alternatives to hunting, photography, and poaching.
In this final chapter, we present the challenges that recent years and especially 2020–21 have presented in the context of trophy hunting.In Chapter 9.1, we cover Covid-19 and how it has affected hunting and conservation differently in Africa, Europe, and the US. African countries and communities have suffered greatly economically from a lack of tourism, which has led to increased poaching and habitat loss; British venison could not be offhanded as it is largely supplied to restaurants, which have been closed because of Covid; and rural hunting and self-sufficiency mentality has increased in the US.In Chapter 9.2, we deal with the increasingly prominent issues of misinformation and disinformation in science communication and communication about trophy hunting especially, and with how social media amplifies misinformation. We describe a couple of the major trophy hunting disinformation campaigns on both sides of the fence and note how both celebrities and certain celebrity scientists seem to use anti trophy hunting campaigning as a popularity booster.
We have three goals in Chapter 4.The first goal is to describe who hunters are (and to a lesser extent antihunters). Demographical information about hunters is much better for USA than for Europe, and trophy hunting is ten times the scale in USA as in the rest of the world combined, so we focus on a portrayal of American hunters. We use mainly the United States Fish and Wildlife Service statistics and the Virginia based Responsive Management survey research firm combined with demographical information about members of the Boone and Crockett Club and Safari Club International. Education, income, gender, ages, race, hunting efforts, and prey species of American hunters is laid out in Chapter 4.1, and we discuss common backgrounds of antihunters in Chapter 4.4.Our second goal (in Chapter 4.2 Hunting, Privilege, and Social Schisms) is to present and defend the hypothesis that hunter-antihunter conflicts are not just about hunting, but about many other social and sociocultural differences and conflicts. The trophy hunter stereotype (based on the demographics described in Chapter 4.1) is a male, white, conservative, protestant, wealthy, pro-gun, business owner. The antihunter (Chapter 4.4) is typically a female, non-white, liberal, anti-gun student. Hunting is just one representation of a mutual dislike that stems from many underlying societal tensions.Our third goal (in Chapters 4.3 and 4.5) is to explore why hunters hunt. We take our departure in the works of Stephen R. Kellert, and supplement with scholars like Jan E. Dizard, Simon Bronner, and Allen Morris Jones to discuss the three different archetypes of hunters (the nature hunter, the meat hunter, and the sport hunter), their reasons for hunting, and what hunting means to them. In Chapter 4.5, we discuss hunting motivations outside or not fully covered by Kellert’s framework and motivations that pertain specifically to trophies.
Anthropomorphism – assigning human characteristics to nonhuman entities – plays an important role in trophy hunting, because the animals that are hunted for trophies are generally some of the most anthropomorphised animals with prominent and often highly anthropomorphized representation in movies. Deer are Bambis, lions are Simbas, elephants are Dumbos, etc. We argue that anthropomorphism is an important reason that trophy hunting is so disdained.We introduce this subject in 7.1 by way of an examination of a giraffe called Marius, who was killed in Copenhagen Zoo in 2014. We use Marius to introduce a discussion of what makes some animals anthropomorphised and loved while others are hated. The various features of animals that research has shown elicit human empathy are covered and the strategies of zoos in using those features for their benefit are explored. We also consider what makes some stories about animals more likely to go viral than others, such as animal names.We discuss what this all means and how it matters in a trophy hunting context in 7.2 and demonstrate that trophy hunted animals have many of the relevant features.
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Measuring rates and causes of mortalities is important in animal ecology and management. Observing the fates of known individuals is a common method of estimating life history variables, including mortality patterns. It has long been assumed that data lost when known animals disappear were unbiased. We test and reject this assumption under conditions common to most, if not all, studies using marked animals. We illustrate the bias for 4 endangered Wolf populations in the United States by reanalyzing data and assumptions about the known and unknown fates of marked wolves to calculate the degree to which risks of different causes of death were mismeasured. We find that, when using traditional methods, the relative risk of mortality from legal killing measured as a proportion of all known fates was overestimated by 0.05-0.16 and the relative risk of poaching was underestimated by 0.17-0.44. We show that published government estimates are affected by these biases and, importantly, are underestimating the risk of poaching. The underestimates have obscured the magnitude of poaching as the major threat to endangered Wolf populations. We offer methods to correct estimates of mortality risk for marked animals of any taxon and describe the conditions under which traditional methods produce more or less bias. We also show how correcting past and future estimates of mortality parameters can address uncertainty about wildlife populations and increase the predictability and sustainability of wildlife management interventions. © The Author 2017. Published by Oxford University Press on behalf of American Society of Mammalogists.
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Trophy hunting has potential to support conservation financing and contribute towards rural development. We conducted a systematic review of the Zimbabwean trophy hunting perspective spanning from pre-1890 to 2015, by examining the following: (1) evolution of legal instruments, administration, and governance of trophy hunting, (2) significance of trophy hunting in conservation financing and rural development, and (3) key challenges, emerging issues in trophy hunting industry, and future interventions. Our review shows that (i) there has been a constant evolution in the policies related to trophy hunting and conservation in Zimbabwe as driven by local and international needs; (ii) trophy hunting providing incentives for wildlife conservation (e.g., law enforcement and habitat protection) and rural communities’ development. Emerging issues that may affect trophy hunting include illegal hunting, inadequate monitoring systems, and hunting bans. We conclude that trophy hunting is still relevant in wildlife conservation and rural communities’ development especially in developing economies where conservation financing is inadequate due to fiscal constraints. We recommend the promotion of net conservation benefits for positive conservation efforts and use of wildlife conservation credits for the opportunity costs associated with reducing trophy hunting off-take levels and promoting non-consumptive wildlife use options.
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Reliable estimates of animal density are fundamental to our understanding of ecological processes and population dynamics. Furthermore, their accuracy is vital to conservation biology since wildlife authorities rely on these figures to make decisions. However, it is notoriously difficult to accurately estimate density for wide-ranging species such as carnivores that occur at low densities. In recent years, significant progress has been made in density estimation of Asian carnivores, but the methods have not been widely adapted to African carnivores. African lions (Panthera leo) provide an excellent example as although abundance indices have been shown to produce poor inferences, they continue to be used to estimate lion density and inform management and policy. In this study we adapt a Bayesian spatially explicit capture-recapture model to estimate lion density in the Maasai Mara National Reserve (MMNR) and surrounding conservancies in Kenya. We utilize sightings data from a three-month survey period to produce statistically rigorous spatial density estimates. Overall posterior mean lion density was estimated to be 16.85 (posterior standard deviation = 1.30) lions over one year of age per 100km(2) with a sex ratio of 2.2♀:1♂. We argue that such methods should be developed, improved and favored over less reliable methods such as track and call-up surveys. We caution against trend analyses based on surveys of differing reliability and call for a unified framework to assess lion numbers across their range in order for better informed management and policy decisions to be made. This article is protected by copyright. All rights reserved.
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There is growing evidence of escalating wildlife losses worldwide. Extreme wildlife losses have recently been documented for large parts of Africa, including western, Central and Eastern Africa. Here, we report extreme declines in wildlife and contemporaneous increase in livestock numbers in Kenya rangelands between 1977 and 2016. Our analysis uses systematic aerial monitoring survey data collected in rangelands that collectively cover 88% of Kenya's land surface. Our results show that wildlife numbers declined on average by 68% between 1977 and 2016. The magnitude of decline varied among species but was most extreme (72-88%) and now severely threatens the population viability and persistence of warthog, lesser kudu, Thomson's gazelle, eland, oryx, topi, hartebeest, impala, Grevy's zebra and waterbuck in Kenya's rangelands. The declines were widespread and occurred in most of the 21 rangeland counties. Likewise to wildlife, cattle numbers decreased (25.2%) but numbers of sheep and goats (76.3%), camels (13.1%) and donkeys (6.7%) evidently increased in the same period. As a result, livestock biomass was 8.1 times greater than that of wildlife in 2011-2013 compared to 3.5 times in 1977-1980. Most of Kenya's wildlife (ca. 30%) occurred in Narok County alone. The proportion of the total "national" wildlife population found in each county increased between 1977 and 2016 substantially only in Taita Taveta and Laikipia but marginally in Garissa and Wajir counties, largely reflecting greater wildlife losses elsewhere. The declines raise very grave concerns about the future of wildlife, the effectiveness of wildlife conservation policies, strategies and practices in Kenya. Causes of the wildlife declines include exponential human population growth, increasing livestock numbers, declining rainfall and a striking rise in temperatures but the fundamental cause seems to be policy, institutional and market failures. Accordingly, we thoroughly evaluate wildlife conservation policy in Kenya. We suggest policy, institutional and management interventions likely to succeed in reducing the declines and restoring rangeland health, most notably through strengthening and investing in community and private wildlife conservancies in the rangelands.
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It is argued that trophy hunting of large, charismatic mammal species can have considerable conservation benefits but only if undertaken sustainably. Social-ecological theory suggests such sustainability only results from developing governance systems that balance financial and biological requirements. Here we use lion (Panthera leo) trophy hunting data from Tanzania to investigate how resource ownership patterns influence hunting revenue and offtake levels. Tanzania contains up to half of the global population of free-ranging lions and is also the main location for lion trophy hunting in Africa. However, there are concerns that current hunting levels are unsustainable. The lion hunting industry in Tanzania is run by the private sector, although the government leases each hunting block to companies, enforces hunting regulation, and allocates them a species-specific annual quota per block. The length of these leases varies and theories surrounding property rights and tenure suggest hunting levels would be less sustainable in blocks experiencing a high turnover of short-term leases. We explored this issue using lion data collected from 1996 to 2008 in the Selous Game Reserve (SGR), the most important trophy hunting destination in Tanzania. We found that blocks in SGR with the highest lion hunting offtake were also those that experienced the steepest declines in trophy offtake. In addition, we found this high hunting offtake and the resultant offtake decline tended to be in blocks under short-term tenure. In contrast, lion hunting levels in blocks under long-term tenure matched more closely the recommended sustainable offtake of 0.92 lions per 1000 km2. However, annual financial returns were higher from blocks under short-term tenure, providing $133 per km2 of government revenue as compared to $62 per km2 from long-term tenure blocks. Our results provide evidence for the importance of property rights in conservation, and support calls for an overhaul of the system in Tanzania by developing competitive market-based approaches for block allocation based on long-term tenure of ten years.
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While trophy hunting provides revenue for conservation, it must be carefully managed to avoid negative population impacts, particularly for long-lived species with low natural mortality rates. Trophy hunting has had negative effects on lion populations throughout Africa, and the species serves as an important case study to consider the balance of costs and benefits, and to consider the effectiveness of alternative strategies to conserve exploited species. Age-restricted harvesting is widely recommended to mitigate negative effects of lion hunting, but this recommendation was based on a population model parameterized with data from a well-protected and growing lion population. Here, we used demographic data from lions subject to more typical conditions, including source-sink dynamics between a protected National Park and adjacent hunting areas in Zambia's Luangwa Valley, to develop a stochastic population projection model and evaluate alternative harvest scenarios. Hunting resulted in population declines over a 25-yr period for all continuous harvest strategies, with large declines for quotas >1 lion/concession (∼0.5 lion/1,000 km²) and hunting of males younger than seven years. A strategy that combined periods of recovery, an age limit of ≥7 yr, and a maximum quota of ∼0.5 lions shot/1,000 km² yielded a risk of extirpation <10%. Our analysis incorporated the effects of human encroachment, poaching, and prey depletion on survival, but assumed that these problems will not increase, which is unlikely. These results suggest conservative management of lion trophy hunting with a combination of regulations. To implement sustainable trophy hunting while maintaining revenue for conservation of hunting areas, our results suggest that hunting fees must increase as a consequence of diminished supply. These findings are broadly applicable to hunted lion populations throughout Africa and to inform global efforts to conserve exploited carnivore populations.
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Effective conservation requires value judgements as well as science (Dickman et al. 2015). The furore over the killing of ‘Cecil’ the lion highlighted the complexities of such judgements. It demonstrated that some people view trophy hunting as morally wrong, and revealed public ignorance that it is a legal, widespread component of African wildlife management, protecting more land than National Parks (Di Minin et al. 2016). Open toleration of trophy hunting by conservationists on these grounds provoked further outrage.
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Many ecological theories and species conservation programmes rely on accurate estimates of population density. Accurate density estimation, especially for species facing rapid declines, requires the application of rigorous field and analytical methods. However, obtaining accurate density estimates of carnivores can be challenging as carnivores naturally exist at relatively low densities and are often elusive and wide-ranging. In this study, we employ an unstructured spatial sampling field design along with a Bayesian sex-specific spatially explicit capture-recapture (SECR) analysis, to provide the first rigorous population density estimates of cheetahs (Acinonyx jubatus) in the Maasai Mara, Kenya. We estimate adult cheetah density to be between 1.28 ± 0.315 and 1.34 ± 0.337 individuals/100km2 across four candidate models specified in our analysis. Our spatially explicit approach revealed ‘hotspots’ of cheetah density, highlighting that cheetah are distributed heterogeneously across the landscape. The SECR models incorporated a movement range parameter which indicated that male cheetah moved four times as much as females, possibly because female movement was restricted by their reproductive status and/or the spatial distribution of prey. We show that SECR can be used for spatially unstructured data to successfully characterise the spatial distribution of a low density species and also estimate population density when sample size is small. Our sampling and modelling framework will help determine spatial and temporal variation in cheetah densities, providing a foundation for their conservation and management. Based on our results we encourage other researchers to adopt a similar approach in estimating densities of individually recognisable species.
Large predators are in decline globally with growing concerns over the impacts of human activity on conservation status and range of many populations. The role of trophy hunting in the conservation or decline of predators is hotly debated, though opposing views are often poorly supported by empirical evidence. Nevertheless an understanding of effects of trophy hunting on populations and behaviour is critical to the conservation of large carnivore populations. The impacts of trophy hunting on African lion population demographics, social structure and spatial behaviour were investigated in Hwange National Park, Zimbabwe, from 1999 to 2012, a period characterized by different trophy hunting intensities. Adult males were primarily targeted by trophy hunters, but survival of all age and sex classes were lowest when male lion off-takes were highest. Reduction in hunting quotas over the study period resulted in a 62% increase in the total population and a 200% increase in adult male density. Adult sex ratios were highly skewed towards females when hunting was intense. Intensity of hunting affected male and female home-range size, which declined in periods of low hunting corresponding to increases in adult males and male coalitions. Trophy hunting on the park boundary exerted a measurable edge effect with lower survival for animals of all age and sex classes living on the park boundary compared to those distant from it. This study provides evidence for negative impacts of uncontrolled trophy hunting on lion population and behaviour. However, limited, well regulated quotas may be compatible with large carnivore conservation.