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BACKGROUND
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,
PERSPECTIVE
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.
Email: david.macdonald@zoo.ox.ac.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.
Email: Andrew.loveridge@zoo.ox.ac.uk
Amy DICKMAN Wildlife Conservation Research Unit, Department of Zoology, The Recanati-Kaplan
Centre, University of Oxford, Tubney House, Tubney, Oxon OX13 5QL, UK.
Email: Amy.dickman@zoo.ox.ac.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.
Email: Paul.johnson@zoo.ox.ac.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.
Email: kimsjacobsen@gmail.com
Byron DU PREEZ Wildlife Conservation Research Unit, Department of Zoology, The Recanati-Kaplan
Centre, University of Oxford, Tubney House, Tubney, Oxon OX13 5QL, UK.
Email: byron.d.dupreez@gmail.com
Mammal Review (2017) © 2017 The Mammal Society and John Wiley & Sons Ltd
Keywords
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
ABSTRACT
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
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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.
2014).
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.
2016).
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
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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
management.
HOW HUNTING AFFECTS LION
POPULATIONS
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
monitoring.
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).
WHAT WOULD HAPPEN TO LION
POPULATIONS IF THEY WERE NO LONGER
HUNTED FOR TROPHIES?
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,
b).
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.
THE CURRENT STATUS OF TROPHY
HUNTING
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.
5
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).
ETHICS AND HINDSIGHT
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.
CONCLUSION
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.
ACKNOWLEDGEMENTS
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
original report.
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