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1
Department
of
Biology,
Colorado
State
University,
1878
Campus
Delivery,
Fort
Collins,
CO
80523,
USA
2
Kellogg
Biological
Station,
Michigan
State
University,
Hickory
Corners,
MI
49060,
USA
3
Graduate
Degree
Program
in
Ecology,
Colorado
State
University,
Fort
Collins,
CO
80523,
USA
*Correspondence:
justin.havird@colostate.edu
(J.C.
Havird).
http://dx.doi.org/10.1016/j.tree.2015.11.012
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Science
&
Society
Banning
Trophy
Hunting
Will
Exacerbate
Biodiversity
Loss
Enrico
Di
Minin,
1,2,
*
Nigel
Leader-Williams,
3
and
Corey
J.A.
Bradshaw
4,5
International
pressure
to
ban
trophy
hunting
is
increasing.
However,
we
argue
that
trophy
hunting
can
be
an
important
conservation
tool,
pro-
vided
it
can
be
done
in
a
controlled
manner
to
benefit
biodiversity
con-
servation
and
local
people.
Where
political
and
governance
structures
are
adequate,
trophy
hunting
can
help
address
the
ongoing
loss
of
species.
International
Outrage
over
Trophy
Hunting
in
Africa
An
American
hunter
killed
a
charismatic
male
lion
(Panthera
leo)
called
Cecil
in
Zimbabwe
in
July
2015.
This
sparked
international
outrage,
mainly
via
a
storm
of
social
and
other
media.
Several
alleged
aspects
of
the
hunt
itself,
such
as
baiting
close
to
national
park
boundaries,
were
done
illegally
and
apparently
against
the
spirit
and
ethical
norms
of
well-managed
trophy
hunts.
Online
outrage
had
also
been
sparked
earlier
in
2015
by
the
legal
hunt
of
a
Critically
Endangered
male
black
rhino
(Diceros
bicornis).
This
hunt
was
sanctioned
by
the
Namibian
Government
via
an
auctioned
permit
that
cost
the
hunter
US$350
000
for
the
privilege.
This
outrage
arose
even
though
the
male
was
considered
‘surplus’
to
the
national
black
rhino
management
plan,
and
the
revenue
generated
from
the
hunt
was
to
be
rein-
vested
into
a
conservation
trust
fund
to
the
wider
good
of
conservation
in
Namibia.
These
two
high-profile
hunts
and
the
ensuing
public
backlash
against
the
ethics
and
conduct
of
trophy
hunting
in
general
have
led
to
proposals
to
ban
the
practice
throughout
Africa.
Furthermore,
some
commercial
passenger
and
cargo
airlines
have
decided
to
stop,
or
may
soon
stop,
the
transport
of
trophies
of
hunted
animals
shot
legally
and
sustainably
by
foreign
tourists,
irrespective
of
international
con-
ventions,
such
as
the
Convention
on
Inter-
national
Trade
in
Endangered
Species
of
Wild
Fauna
and
Flora
(CITES)
and
national
laws
that
allow
trophy
hunting.
Hunting
Industry
in
Sub-Saharan
Africa
Trophy
hunting
strongly
contributes
to
the
conservation
enterprise
in
sub-Saharan
Africa,
where
large
areas
support
important
terrestrial
biodiversity
that
is
currently
allo-
cated
to
trophy
hunting
use
(Table
1).
While
most
of
the
hunted
individuals
(e.g.,
96%
in
South
Africa
in
2012)
[1]
are
often
from
more
common
and
less
valuable
species
(Table
1),
most
of
the
trophy
hunting
reve-
nue
is
generated
from
a
few
species
carry-
ing
valuable
trophies,
particularly
the
charismatic
‘Big
Five’
(lion
leopard
Pan-
thera
pardus;
elephant
Loxodonta
africana;
buffalo
Syncerus
caffer;
and
black
or
white
rhinoceros
Ceratotherium
simum)
[2].
Out
of
the
US$68
million
of
gross
revenue
gen-
erated
from
trophy
hunting
in
South
Africa
in
2012,
over
US$28
million
(at
least
41%)
was
generated
from
the
Big
Five
alone
(i.e.,
$5
635
625
from
635
buffaloes;
$1
194
600
from
33
elephants;
$647
500
from
37
leopards;
$15
270
750
from
617
lions,
$300
000
from
one
black
rhinoceros;
and
$5
355
000
from
63
white
rhinoceroses)
[1].
Southern
African
countries
and
Tanza-
nia
exported
most
of
the
Big
Five
trophies
between
2009
and
2013
(Figure
1).
At
the
same
time,
two
countries
that
do
not
typi-
cally
attract
many
tourists
(the
Central
Afri-
can
Republic,
currently
undergoing
a
conflict,
and
Cameroon,
where
poaching
pressure
is
high)
allowed
trophy
hunting
of
big
cats
and
elephants,
respectively,
over
the
same
period
(Figure
1).
Concerns
about
Trophy
Hunting
Overall,
land
allocated
to
trophy
hunting
has
the
potential
to
assist
countries
to
achieve
biodiversity
conservation
goals
[3].
However,
the
contribution
of
hunting
to
conservation
is
often
contentious
for
various
reasons.
There
can
be
uncertainty
over
the
sustainability
of
offtake
rates
and
their
potential
impact
on
wildlife
popula-
tions
[4].
This
concern
arises
because
quotas
and
offtakes
are
not
often
based
on
scientific
assessments.
Furthermore,
restrictions
on
the
age
of
hunted
Trends
in
Ecology
&
Evolution,
February
2016,
Vol.
31,
No.
2
99
individuals
are
not
often
implemented
[5].
In
addition,
the
contribution
of
some
forms
of
trophy
hunting
to
conservation
is
debat-
able.
This
is
particularly
the
case
for
‘canned
lion
hunting’,
where
future
targets
are
bred
and
raised
in
captivity
and
kept
in
confined
enclosures
until
shot,
to
ensure
that
hunters
are
guaranteed
a
kill.
In
South
Africa,
which
is
by
far
the
largest
exporter
of
lion
trophies
across
sub-Saharan
Africa
(Figure
1),
80%
of
the
trophies
between
2009
and
2013
were
from
lions
raised
in
captivity
or
ranched.
The
ethics
of
canned
hunting
are
dubious,
and
this
abhorrent
practice
requires
reform
before
it
brings
down
ethically
practiced
hunting.
The
profitability
of
their
respective
hunting
industries
is
hard
to
compare
across
sub-
Saharan
countries
[5].
Nevertheless,
it
is
known
that
the
gross
annual
revenue
gen-
erated
by
the
hunting
industry
comprises
tens
of
millions
of
US$
in
countries
such
as
South
Africa,
Tanzania,
and
Botswana
(Table
1).
Despite
this,
the
amount
of
accrued
revenue
allocated
to
conserva-
tion
authorities
that
could
in
principle
be
reinvested
in
improved
management
appears
to
be
limited.
In
Tanzania,
for
example,
the
accrued
revenue
allocated
to
the
Wildlife
Division
in
2008
amounted
to
22%
(US$12
353
180)
of
the
gross
revenue
generated
by
hunting
in
that
year
[5].
The
remainder
of
the
revenue
went
to
the
private
sector.
Another
limitation
is
that
revenue
gener-
ated
from
trophy
hunting
currently
pro-
vides
few
benefits
to
local
communities
sharing
habitats
with
biodiversity
[6,7].
In
Namibia,
however,
revenue
generated
from
trophy
hunting
has
encouraged
local
community
participation
in
conser-
vation,
which
in
turn
has
resulted
in
sub-
stantial
increases
in
the
abundance
of
many
wildlife
species
and
in
the
total
area
of
land
falling
under
community
protection
through
conservancies
[6].
It
is
less
clear
in
other
African
countries
what
proportions
of
hunting-permit
rev-
enue
are
directed
to
community-devel-
opment
projects,
whether
they
are
payments
to
community-based
orga-
nizations,
or
payments
to
communities
for
concession
fees,
resource
fees,
or
payments
for
welfare
and
education.
Finally,
legal
controls
over
biological,
eth-
ical,
and
financial
aspects
of
the
hunting
industry
can
be
more
easily
circum-
vented
in
many
sub-Saharan
countries
where
management
capacity
and
gover-
nance
structures
are
ineffective
[8].
Why
Blanket
Bans
Could
Exacerbate
Biodiversity
Loss
One
currently
promoted
solution
to
address
such
concerns
is
to
ban
trophy
hunting
altogether.
However,
a
blanket
ban
on
trophy
hunting
could
lead
to
worse
conservation
outcomes
for
three
main
reasons.
First,
financial
resources
for
conservation
are
limited,
particularly
in
developing
countries.
Hence,
both
nonconsumptive
and
consumptive
uses
of
wildlife
are
necessary
to
generate
enough
funding
to
support
meaningful
conservation
success
over
large
areas
[9].
While
ecotourism
can
help
reduce
poverty
in
communities
coexisting
with
biodiversity
[3],
ecotourists
generally
pre-
fer
travelling
to
more
accessible
areas
[10],
greatly
limiting
the
opportunities
for
con-
servation
in
more
remote
regions.
Instead,
sustainable
hunting
can
create
important
incentives
for
biodiversity
conservation
in
areas
where
ecotourism
is
not
economi-
cally
viable
[11].
At
a
time
when
greater
proportions
of
conservation
budgets
are
being
spent
on
enforcement,
the
revenue
from
trophy
hunting
can
empower
Table
1.
Hunting
Contribution
to
Biodiversity
Conservation
and
National
Economies
in
Sub-Saharan
Countries
Country
Area
Covered
by
Game
Ranches
(%
of
Total
Land
Area)
a
Terrestrial
Protected
Areas
(%
of
Total
Land
Area)
b
Top
3
Most
Exported
Trophies
in
2012
c,d
Annual
Revenue
(US$
million)
e
South
Africa
13.1
6.2
impala,
warthog,
kudu
68.0
f
Tanzania
26.4
32.2
leopard,
hippopotamus,
elephant
56.3
g
Botswana
23.0
37.2
elephant,
leopard,
lechwe
40.0
g
Namibia
11.4
43.2
zebra,
chacma
baboon,
leopard
28.5
h
Zimbabwe
16.6
27.2
elephant,
leopard,
chacma
baboon
15.8
g
Mozambique
10.5
17.6
Nile
crocodile,
elephant,
hippopotamus
5.0
g
Zambia
21.3
37.8
lechwe,
hippopotamus,
leopard
3.6
g
Total
–
–
–
217.2
a
[13].
b
World
Bank
(http://data.worldbank.org/indicator/ER.LND.PTLD.ZS).
c
CITES
trade
database
(http://trade.cites.org/).
d
Chacma
baboon
(Papio
ursinus),
elephant
(Loxodonto
africana),
hippopotamus
(Hippopotamus
amphibius),
impala
(Aepyceros
melampus),
greater
kudu
(Tragelaphus
strepsiceros),
lechwe
(Kobus
leche),
leopard
(Panthera
pardus),
Nile
crocodile
(Crocodylus
niloticus),
warthog
(Phacochoerus
africanus),
zebra
(Equus
quagga).
e
Data
not
adjusted
for
inflation.
f
[1].
g
[6]
(data
for
2008
for
Botswana,
Tanzania
and
Mozambique,
2007
for
Zimbabwe,
and
2002
for
Zambia).
h
African
Indaba
(http://www.africanindaba.com/wp-content/uploads/2014/03/AfricanIndabaVol3-4.pdf).
100
Trends
in
Ecology
&
Evolution,
February
2016,
Vol.
31,
No.
2
communities
to
protect
their
resources
by
the
employment
of
more
antipoaching
rangers
or
the
construction
of
disincentive
infrastructure
[12].
If
revenue
cannot
be
generated
from
trophy
hunting,
natural
habitats
will
be
transformed
to
other
forms
of
land
use
that
provide
higher
return
on
investments
compared
with
conservation
[3],
but
have
negative
impacts
on
biodiversity.
Second,
trophy
hunting
can
have
a
smaller
footprint
than
ecotourism
in
terms
of
carbon
emissions,
infrastructure
devel-
opment,
and
personnel,
and
can
generate
more
revenue
from
a
lower
volume
of
tourist
hunters.
An
often-neglected
rela-
tion
exists
between
ecotourism
and
avia-
tion
with
regard
to
energy
use
and
greenhouse
gas
emissions.
Compared
with
ecotourism,
the
trophy-hunting
industry
relies
on
fewer
tourist
hunters,
because
the
income
generated
per
hunter
is
higher
[13].
Additionally,
hunters
are
interested
in
maintaining
good-quality
habitat
for
the
simple
reason
that
the
qual-
ity
of
the
individuals
harvested
therein
is
also
high
[14].
Finally,
hunters
are
pre-
pared
to
hunt
in
areas
lacking
attractive
scenery,
and
require
less
infrastructure,
therefore
minimizing
habitat
degradation.
Third,
management
for
hunting
places
emphasis
on
maintaining
large
wildlife
populations
for
offtake,
as
opposed
to
ecotourism,
where
the
presence
of
only
a
few
individual
animals
is
sufficient
to
maximize
profits
[2].
Both
the
consump-
tive
and
nonconsumptive
uses
of
biodiver-
sity
can
generate
important
revenue,
so
allowing
local
stakeholders,
such
as
pri-
vate
landowners
and
communities,
to
retain
property
rights
over
these
species
is
a
necessary
precursor
for
them
to
justify
offsetting
the
direct
and
opportunity
costs
of
conservation.
Thus,
the
economic
models
underlying
ecotourism
and
trophy
hunting
may
lead
to
diverging
manage-
ment
strategies.
Empirical
evidence
shows
that
the
strategy
of
artificially
man-
aging
small
populations
within
electrified
fences
to
maximize
economic
return
from
Black rhino (Diceros bicomis)
Crically endangered
CITES appendix I
Populaon trend increasing
Leopard (Panthera pardus)
Near threatened
CITES appendix I
Populaon trend decreasing
8
4
2
343
200
100
Elephant (Loxodonta africana)
Vulnerable
CITES appendix I and II
Populaon trend increasing
496
400
200
Lion (Panthera leo)
Vulnerable
CITES appendix II
Populaon trend decreasing
877
600
300
Hippo (Hippopotamus amphibius)
Vulnerable
CITES appendix II
Populaon trend decreasing
White rhino (Ceratotherium simum)
Near threatened
CITES appendix I and II
Populaon trend increasing
328
200
100
191
100
50
Figure
1.
Number
of
Trophies
Exported
from
2009
to
2013
(Red
Bars)
for
Six
Charismatic
African
Species
Subject
to
Trophy
Hunting.
Gray-shaded
areas
correspond
to
the
range
maps
of
species
obtained
from
www.iucnredlist.org/technical-documents/spatial-data.
Each
species
is
listed
under
the
Convention
on
International
Trade
in
Endangered
Species
of
Wild
Fauna
and
Flora
(CITES).
Net
export
data
for
each
species
retrieved
from
the
CITES
Trade
Database
(http://trade.cites.org)
by
searching
for
‘trophies’
under
the
‘trade
terms’
bar.
Information
about
the
conservation
status,
population
trend,
and
CITES
listing
retrieved
from
www.
iucnredlist.org
and
www.cites.org.
Numbers
next
to
the
external
bar
in
each
panel
indicate
the
scale
to
interpret
bar
charts
of
annual
trophies
taken
per
species
and
per
country.
Trends
in
Ecology
&
Evolution,
February
2016,
Vol.
31,
No.
2
101
ecotourism
and
minimize
management
costs
might
be
the
most
appropriate
to
enhance
tourist
experiences
[2].
By
con-
trast,
wildlife
populations
potentially
have
higher
hunting
value
when
their
sizes
are
larger
(i.e.,
are
more
viable)
and
popula-
tions
are
better
connected
to
enhance
gene
flow,
because
the
latter
can
affect
the
fitness
and
quality
of
the
individuals
harvested.
Concluding
Remarks
Inadequate
political,
legal,
and
governance
structures
are
currently
preventing
trophy
hunting
from
being
an
effective
tool
for
cre-
ating
conservation
incentives
in
sub-
Saharan
Africa.
At
the
same
time,
banning
trophy
hunting
might
not
be
the
best
solu-
tion
because
biodiversity
loss
could
even
be
worse
in
its
absence.
Therefore,
we
propose
a
set
of
prescriptions
that
could
enhance
the
contribution
of
trophy
hunting
to
conservation
and
to
the
equitable
shar-
ing
of
the
benefits
with
local
people
(Box
1).
To
make
these
prescriptions
more
relevant
for
decision-makers,
we
have
summarized
them
according
to
the
guiding
principles
on
trophy
hunting
promoted
by
the
Interna-
tional
Union
for
the
Conservation
of
Nature
[15].
In
particular,
we
make
suggestions
on
how
net
biodiversity
benefits
and
stake-
holder
returns
can
be
achieved
simulta-
neously,
and
highlight
how
the
hunting
industry
and
governance
structures
can
be
made
more
transparent
to
avoid
unethi-
cal
or
illegal
practices.
Finally,
we
provide
additional
guidelines
to
account
for
animal
welfare
concerns.
Promoting
these
and
other
prescriptions
could
enhance
the
role
of
trophy
hunting
in
addressing
the
ongoing
loss
of
species.
Acknowledgments
We
thank
M.
Festa-Bianchet
and
an
anonymous
reviewer
for
comments
that
helped
improve
the
man-
uscript.
E.D.M.
acknowledges
ERC-StG
Grant
260393
(GEDA)
and
the
Academy
of
Finland
Centre
of
Excellence
Programme
2012–2017
(Grant
250444)
for
support.
Unless
spec-
ified
otherwise,
photographs
in
Figure
1
by
E.D.M.
1
Department
of
Biosciences,
University
of
Helsinki,
FI-00014,
Helsinki,
Finland
2
School
of
Life
Sciences,
University
of
KwaZulu-Natal,
Private
Bag
X
54001,
Durban
4000,
South
Africa
3
Department
of
Geography,
University
of
Cambridge,
Downing
Place,
Cambridge,
CB2
3EN,
UK
4
The
Environment
Institute
and
School
of
Biological
Sciences,
The
University
of
Adelaide,
Adelaide,
SA
5005,
Australia
5
Laboratoire
Écologie,
Systématique
et
Évolution
–
UMR
CNRS
8079,
Université
Paris-Sud,
Bat
362,
F-91405,
Orsay
Cedex,
France
*Correspondence:
enrico.di.minin@helsinki.fi
(E.
Di
Minin).
http://dx.doi.org/10.1016/j.tree.2015.12.006
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Box
1.
Prescriptions
to
Make
Trophy
Hunting
More
Effective
for
Conservation
1.
Net
Conservation
Benefit
1.1
Mandatory
levies
imposed
on
safari
operators
by
governments
that
are
invested
directly
into
trust
funds
for
conservation
and
management.
1.2
Ecolabeling
certification
schemes
adopted
for
trophies
originating
from
areas
that
contribute
to
broader
biodiversity
conservation
and
respect
animal
welfare
concerns.
2.
Biological
Sustainability
2.1
Mandatory
population
viability
analyses
to
ensure
that
harvests
cause
no
net
population
decline.
2.2
Ban
posthunt
sales
of
any
portion
of
the
quarry
shot,
to
avoid
illegal
wildlife
trade.
3.
Socio-Economic-Cultural
Benefit
3.1
Promote
and
fund
trophy-hunting
enterprises
run
(or
leased)
by
local
communities.
3.2
Create
trusts
to
facilitate
equitable
benefit
sharing
within
local
communities
and
promote
long-term
economic
sustainability.
4.
Adaptive
Management:
Planning,
Monitoring,
and
Reporting
4.1
Mandatory
scientific
sampling
of
hunted
animals,
including
tissue
for
genetics,
teeth
for
age
analysis,
stomach
contents,
full
morphometrics,
disease
screening,
and
so
on.
4.2
Mandatory
5-year
reviews
of
all
individuals
hunted
and
detailed
population
management
plans
submitted
to
government
legislators
to
extend
permits.
5.
Accountable
and
Effective
Governance
5.1
Full
disclosure
to
the
public
of
all
data
collected
(including
levied
amounts),
although
personal
details
of
proponents
held
by
government
legislators
only.
5.2
Independent
government
observers
placed
randomly
and
without
forewarning
on
safari
hunts
as
they
happen.
5.3
Trophies
are
confiscated
and
permits
are
revoked
when
illegal
practices
are
disclosed.
6.
Animal
welfare
6.1
Backup
professional
shooters
and
trackers
present
for
all
hunts
to
minimize
welfare
concerns
102
Trends
in
Ecology
&
Evolution,
February
2016,
Vol.
31,
No.
2