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International law and lions (Panthera leo): Understanding and improving the contribution of wildlife treaties to the conservation and sustainable use of an iconic carnivore

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The lion (Panthera leo) is featuring ever more prominently on the agendas of international wildlife treaties like the Convention on International Trade in Endangered Species (CITES) and the Convention on the Conservation of Migratory Species of Wild Animals (CMS). Lion range and numbers have declined markedly over the last two decades. In this review we assess the present role of international wildlife treaties with a view to improving their combined contribution to the conservation and sustainable use of lions. Our analysis identifies a substantial body of relevant international wildlife law and, moreover, a significant potential for enhancing the contribution to lion conservation of these global and regional treaties. The time is right to invest in such improvements, and our review renders a range of general and treaty-specific recommendations for doing so, including making full use of the Ramsar Wetlands Convention, World Heritage Convention and transboundary conservation area (TFCA) treaties for lion conservation. The CMS holds particular potential in this regard and our analysis provides strong support for listing the lion in its Appendices.
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International law and lions (Panthera leo): understanding and improving the contribution... 83
International law and lions (Panthera leo):
understanding and improving the contribution
of wildlife treaties to the conservation and
sustainable use of an iconic carnivore
Arie Trouwborst1, Melissa Lewis1, Dawn Burnham2, Amy Dickman2, Amy Hinks2,
Timothy Hodgetts2, Ewan A. Macdonald2, David W. Macdonald2
1 Department of European and International Public Law, Tilburg University, Tilburg, e Netherlands
2Wildlife Conservation Research Unit (WildCRU), University of Oxford, Tubney, United Kingdom
Corresponding author: Arie Trouwborst (a.trouwborst@tilburguniversity.edu)
Academic editor: S. Williams|Received10 June 207|Accepted 8 August 2017|Published 13 September 2017
http://zoobank.org/B0E33B35-52C0-45B9-8A54-4A60F6D1FE5C
Citation: Trouwborst A, Lewis M, Burnham D, Dickman A, Hinks A, Hodgetts T, Macdonald EA, Macdonald DW
(2017) International law and lions (Panthera leo): understanding and improving the contribution of wildlife treaties to
the conservation and sustainable use of an iconic carnivore. Nature Conservation 21: 83–128. https://doi.org/10.3897/
natureconservation.21.13690
Abstract
e lion (Panthera leo) is featuring ever more prominently on the agendas of international wildlife treaties
like the Convention on International Trade in Endangered Species (CITES) and the Convention on the
Conservation of Migratory Species of Wild Animals (CMS). Lion range and numbers have declined mark-
edly over the last two decades. In this review we assess the present role of international wildlife treaties with a
view to improving their combined contribution to the conservation and sustainable use of lions. Our analy-
sis identies a substantial body of relevant international wildlife law and, moreover, a signicant potential
for enhancing the contribution to lion conservation of these global and regional treaties. e time is right
to invest in such improvements, and our review renders a range of general and treaty-specic recommenda-
tions for doing so, including making full use of the Ramsar Wetlands Convention, World Heritage Conven-
tion and transboundary conservation area (TFCA) treaties for lion conservation. e CMS holds particular
potential in this regard and our analysis provides strong support for listing the lion in its Appendices.
Keywords
Lion, law, international wildlife law, Convention on Migratory Species (CMS), Convention on Interna-
tional Trade in Endangered Species (CITES), Ramsar Wetlands Convention, World Heritage Conven-
tion, Convention on Biological Diversity (CBD)
Nature Conservation 21: 83–128 (2017)
doi: 10.3897/natureconservation.21.13690
http://natureconservation.pensoft.net
Copyright Arie Trouwborst et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
REVIEW ARTICLE
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Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
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Introduction
Lion (Panthera leo) conservation features prominently on the agendas of interna-
tional wildlife treaties like the 1973 Convention on International Trade in Endan-
gered Species of Wild Fauna and Flora (CITES) and the 1979 Convention on the
Conservation of Migratory Species of Wild Animals (CMS or Bonn Convention).
Lion range and numbers have declined markedly over the last two decades (Bauer
et al. 2016). In this review we assess the present role of international wildlife trea-
ties with a view to improving their combined contribution to the conservation and
sustainable use of the lion.
International law and large carnivores
Within the broad arena of the ongoing global biodiversity crisis (Ceballos et al. 2015),
large-bodied species are generally more vulnerable than small-bodied species, and their
population trends reect this (Di Marco et al. 2014; Ripple et al. 2014; Ripple etal.
2015). With some exceptions, such as most European large carnivore populations
(Chapron et al. 2014), the world’s largest carnivores, including lions, are declining,
with range contractions and worsening conservation status (Ripple et al. 2014; Bauer
et al. 2016). Given the important ecological roles of large carnivores, their demise
tends to have negative ecological impacts for other species and ecosystems too (Ripple
et al. 2014). Recently, a large number of conservation scientists involved with large
carnivore and large herbivore conservation called for ‘comprehensive actions to save
these iconic wildlife species’, appealing to all disciplines involved, and duly noting the
role of international wildlife conservation treaties as part of this joint endeavor (Ripple
et al. 2016a).
In the overall eort to stem and reverse biodiversity loss, law is a crucial instru-
ment (Chapron et al. 2017), including international wildlife law (Bowman et al.
2010; Trouwborst et al. 2017c). International wildlife law – alternatively referred to
as international nature conservation law or international biodiversity law – consists
mainly of intergovernmental agreements aimed at conservation of (terrestrial and ma-
rine) species, natural areas, ecosystems, and/or biodiversity at large. ese have been
adopted by states, inter alia, with a view to the transboundary movements and occur-
rence of wildlife populations; the international nature of some of the threats to wild-
life; and the notion that biodiversity conservation is considered a ‘common concern
of mankind’, as recorded in the preamble to the 1992 Convention on Biological Di-
versity (CBD). Eective conservation calls for cross-border approaches and long-term
commitments. International law is the pre-eminent mechanism for realizing these,
and despite the inherent limitations of international treaties and the various chal-
lenges to their eective implementation, many species would have been (even) worse
o without international wildlife law (Bowman et al. 2010; Gillespie 2011; Bowman
2016; Trouwborst et al. 2017c).
International law and lions (Panthera leo): understanding and improving the contribution... 85
International wildlife treaties have contributed to biodiversity conservation in many
dierent ways, including through protected areas designated pursuant to international
commitments; similarly instigated national legislation regulating wildlife exploitation;
enhanced priority accorded to conservation issues on governments’ agendas; incorpo-
ration of technical guidance adopted by treaty bodies into national action plans and
legislation; coordinated collection of data; increased cooperation among and between
governmental and non-governmental stakeholders; direct assistance to conservation
initiatives through treaties’ funding mechanisms; and through many instances where
harmful developments were blocked or particular conservation actions taken when gov-
ernments were confronted with their international obligations in (inter)national court
proceedings or compliance mechanisms (Bowman et al. 2010; Gillespie 2011; Trouw-
borst 2015a; Bowman 2016; Scott 2016; Trouwborst et al. 2017c). ere still appears to
be signicant room for increasing the contribution made by international wildlife law to
conservation, not only by enhancing the legal framework itself, but also by maximising
the legal instruments currently available (Trouwborst 2015a; Bowman 2016).
Across the globe, large carnivores present a special set of conservation issues from
a legal perspective, given inter alia their great spatial requirements, elevated human-
wildlife conict potential, and roles as keystone and/or umbrella species (Macdonald
et al. 2013; Trouwborst 2015a; Treves et al. 2015). For these reasons, and because
of the transboundary nature of many large carnivore populations and some of their
threats, international law has a distinct role (Trouwborst 2015a), though this has re-
ceived little attention in the scholarly literature. Most in-depth research on interna-
tional law and large carnivores has focused on wolves (Canis lupus), brown bears (Ursus
arctos) and lynx (Lynx lynx) in Europe (for a range of examples, see www.clawsandlaws.
eu and www.tilburguniversity.edu/iuscarnivoris), with only one general review of the
relevance of international wildlife law for the world’s 31 largest terrestrial carnivores
(Trouwborst 2015a), and one initial analysis focusing on lions in Africa (Watts 2016).
Lions and international law
e lion is archetypal in all of the aforementioned respects. Given its ecological impor-
tance as an apex predator, it is a keystone species. It is also an umbrella species, in that
lion conservation tends to benet a range of other species (Caro 2003; Macdonald et
al. 2012; Dickman et al. 2015). Lions certainly have large spatial requirements, and
coexistence with humans, particularly outside protected areas, is often problematic
(Loveridge et al. 2010). ere is, moreover, a strong international dimension to lion
conservation. Many of the currently remaining lion populations straddle international
boundaries (Dickman et al. submitted); close links exist between the conservation and
management of lions and international tourism and trophy hunting; the recognition
of many natural areas in Africa as sites of international importance under conservation
treaties is intimately linked to the presence of lions (Watts 2016); there is an increas-
ing international trade in lion parts (Williams et al. 2015; Williams et al. submitted a;
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
86
Williams et al. submitted b); and the worrying conservation status of lions is of inter-
national concern to conservationists and to the global public (Macdonald et al. 2016)
as one of the most iconic and charismatic species (Macdonald et al. 2015).
Globally, the lion has featured on the IUCN Red List as ‘Vulnerable’ since 1996.
Numbers of wild lions have been steadily decreasing and the global population may
be approaching 20,000, with the species persisting in only 8–17% of its historic
range (Riggio et al. 2013; Bauer et al. 2016; Dickman et al. submitted). According
to the latest Red List assessment, lions remain in 25 sub-Saharan African countries
and in a small part of India (Bauer et al. 2016). ey have gone extinct in 26 African
and Eurasian countries; and are ‘possibly extinct’ in 7 African countries (Bauer et al.
2016). Dickman et al (submitted) have recently mapped the 60 known remaining
populations of lions, and only six of these populations consist of more than 1,000
individuals: Selous-Niassa, Serengeti-Mara, Kavango-Zambezi, Greater Limpopo,
Katavi-Ruaha and Kgalagadi (see Figure 1). Just under half of the wild lion estate
lies within protected areas, and Lindsey et al. (2017) have demonstrated that even
there, in most cases the lions are thought to live well below carrying capacity and
at considerable threat from infra-structural inadequacies largely derived from short-
age of funds. ere is a marked dierence between the sharp declines observed in
most range states, and the situation in four southern African countries (Botswana,
Namibia, South Africa, Zimbabwe) and India, where lion populations have declined
only slightly, or are stable or increasing (Bauer et al. 2016). e West African lion
subpopulation is listed as ‘Critically Endangered’ (Henschel et al. 2015). e only
remaining population of Asiatic lions (Panthera leo persica) is considered ‘Endan-
gered’ (Breitenmoser et al. 2008), although local human attitudes have been remark-
ably benign (Venkataraman et al. 2014).
reats to lions include direct persecution, mainly retaliatory or preventive kill-
ing to protect livestock or human life; the depletion of their prey base, mainly due to
poaching in connection with an unsustainable bushmeat trade (see also Ripple et al.
2016b; Sandom et al. 2017); habitat loss; and killing fueled by an increasing demand
for lion bones and body parts (Bauer et al. 2016; Panthera et al. 2017). e rst two
of these threats – human-lion conict and bushmeat poaching – are considered the
gravest (Panthera et al. 2017). Trophy hunting can have positive or negative impacts,
depending on how well it is regulated (Bauer et al. 2016; Loveridge et al. 2016; Mac-
donald 2016; Macdonald et al. 2017).
Dickman et al’s (submitted) rangewide analysis identies for each population, and
for each country within which lions still occur, the intersection of ecological and infra-
structural fragilities. e latter forms a backcloth against which to consider the pattern
of international law in those same countries. Against this backcloth, and building on
Watts (2016), this review aims to explore the current and potential future utility of
international wildlife law for lion conservation. Experience, including our own, indi-
cates that this is best achieved through a multidisciplinary approach (Macdonald and
Chapron 2017), whereby legal experts join forces with ecologists and experts from
other disciplines with a good understanding of the broader context and the actual
International law and lions (Panthera leo): understanding and improving the contribution... 87
Figure 1. Extant lion range (excluding small fenced reserves), Ramsar-listed sites and World Heritage
sites. e numbers indicate the locations of the sites listed in Tables 3 and 4.
conservation needs of species. Such cooperation has, encouragingly, been gathering
momentum in recent years (Cliquet et al. 2009; Trouwborst et al. 2015; Epstein et al.
2016; Selier et al. 2016; Treves et al. 2017; Trouwborst et al. 2017a; Chapron et al.
2017; Redpath et al. 2017; Trouwborst et al. 2017c). Our review, performed by legal
experts, conservation biologists and social scientists, builds on this momentum.
ough focus is thus on lions, the results of our review are likely to be relevant also
for other large carnivore species, particularly in Africa, such as leopard (Panthera par-
dus), cheetah (Acinonyx jubatus), wild dog (Lycaon pictus) and hyaenas (Crocuta crocuta,
Hyaena hyaena, Hyaena brunnea).
Method
Our analysis is based on standard legal research methodology, involving the selection
and interpretation of international legal instruments of relevance to lion conserva-
tion (Trouwborst 2015b). For reasons of space, we limit this analysis to international
wildlife law, although we note the existence of other elds of international law with
direct or indirect signicance for lion conservation, such as legal instruments dealing
with crime, corruption, climate change, or indeed the regulation of pesticides, some
of which are used to poison lions (Watts 2016). For each legal instrument, we oer a
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
88
concise explanation of the most relevant legal obligations (for more exhaustive infor-
mation on those obligations and general background concerning the treaty regimes
involved we refer readers to works such as Bowman et al. (2010) and Gillespie (2011),
and the websites of the various treaties). On that basis, we analyze the various legal
instruments and obligations within their broader context, incorporating knowledge
and insights regarding lions and their conservation needs, and regarding the varying,
real-world concerns of the various lion range states and their human populations. We
focus on the 33 lion range states identied in the IUCN Red List assessment, including
7 states where lions are considered ‘possibly extinct’. We do so in particular with a view
to the potential for lion recolonization or reintroduction.
Overview of the international law and policy framework for lion con-
servation
Binding instruments
Treaties of importance to lion conservation are listed in Table 1. Table 2 and Figure 2
indicate the extent to which the various lion range states are currently bound by eight
of these lion-related treaties under international law as contracting parties. e meth-
ods employed by the various treaties vary. Some treaties operate on the basis of species
lists, with a particular legal regime associated with each list; others involve the listing of
sites; yet others do not employ lists. e treaties’ geographic scopes also vary.
Five treaties are global. ese ‘Big 5’ of international wildlife law are the 1971
Convention on Wetlands of International Importance Especially as Waterfowl Habitat
(Ramsar Convention), the 1972 UNESCO Convention Concerning the Protection
of the World Cultural and Natural Heritage (World Heritage Convention or WHC),
CITES, CMS and CBD. Lions have been in the spotlight mostly in connection with
CITES and, in recent years, the CMS. In May 2016, CITES and CMS jointly hosted
an intergovernmental meeting in Entebbe, which was dedicated specically to African
Table 1. Treaties of relevance to lion conservation. e relevance of each treaty or category of treaties is
indicated for African lion subpopulations and Asiatic lion, respectively. N/A = not applicable.
African lion Asiatic lion (P. leo persica)
Ramsar Convention Habitat in 39 listed sites No listed habitat
World Heritage Convention Habitat in 18 listed sites No listed habitat
CITES Listed in Appendix II Listed in Appendix I
CMS Not (yet) listed, but covered Not (yet) listed, but covered
CBD General relevance General relevance
African Convention Listed in Annex, Class B N/A
Bern Convention Not listed, but covered N/A
SADC Protocol General relevance N/A
Lusaka Agreement General relevance N/A
TFCA treaties General relevance N/A
International law and lions (Panthera leo): understanding and improving the contribution... 89
Table 2. Lion range states and their participation in relevant treaties. List of lion range states as provided
in the IUCN Red List of reatened Species 2016 (excluding previous range states in which the species
is known to be extinct), indicating their participation in relevant treaties. PE = possibly extinct; X = con-
tracting party; - = not currently a contracting party, but could become one; N/A = not applicable (i.e. the
country falls outside of the instrument’s geographic scope).
Range state Ramsar WHC CITES CMS CBD African
Convention
SADC
Protocol
Lusaka
Agreement
Angola - X X X X - - -
Benin X X X X X - N/A -
Botswana X X X - X - X -
Burkina Faso X X X X X X N/A -
Cameroon X X X - X X N/A -
Central African
Republic X X X - X X N/A -
Chad X X X X X - N/A -
Côte d’Ivoire (PE) X X X X X X N/A -
Dem. Rep. of
Congo X X X X X X - -
Ethiopia - X X X X - N/A -
Ghana (PE) X X X X X X N/A -
Guinea (PE) X X X X X X N/A -
Guinea-Bissau (PE) X X X X X - N/A -
India X X X X X N/A N/A N/A
Kenya X X X X X X N/A X
Malawi X X X - X X X -
Mali (PE) X X X X X X N/A -
Mozambique X X X X X X X -
Namibia X X X - X - X -
Niger X X X X X X N/A -
Nigeria X X X X X X N/A -
Rwanda (PE) X X X X X X N/A -
Senegal X X X X X X N/A -
Somalia - - X X X - N/A -
South Africa X X X X X - X -
South Sudan X X - - X - N/A -
Sudan X X X - X X N/A -
Swaziland X X X X X X - -
Togo (PE) X X X X X X N/A -
Uganda X X X X X X N/A X
Un. Rep. of
Tanzania X X X X X X X X
Zambia X X X - X X X X
Zimbabwe X X X X X - X -
33 30 32 32 25 33 21 8 4
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
90
Figure 2. e map shows to how many of the 8 lion-related treaties mentioned in Table 2 each lion range
state is a contracting party.
lion conservation and was attended by delegations of 28 of the 33 range states. e ve
global treaties are analyzed separately below.
Relevant regional treaties are the 1968 African Convention on the Conservation
of Nature and Natural Resources (African Convention), the 1994 Agreement on Co-
operative Enforcement Operations Directed at Illegal Trade in Wild Fauna and Flora
(Lusaka Agreement), the 1999 Protocol (to the 1992 Treaty of the Southern African
Development Community) on Wildlife Conservation and Law Enforcement (SADC
Protocol), and various treaties establishing transfrontier conservation areas (TFCAs).
Curiously, even the 1979 Convention on the Conservation of European Wildlife and
Natural Habitats (Bern Convention) is of potential, albeit more marginal, signicance
to lion conservation (see below). Pertinent instruments that have not yet entered into
force include the 2003 revision of the African Convention and the 2005 Protocol on
Environment and Natural Resources Management to the 1999 Treaty for the Estab-
lishment of the East African Community (EAC Treaty) – although we note the rel-
evance to lion conservation of some provisions of the EAC Treaty itself.
Below, we provide individual analyses of the most relevant treaties, in particular
the Big 5 global conventions, in chronological order of their adoption, followed by
selected regional instruments.
International law and lions (Panthera leo): understanding and improving the contribution... 91
Non-binding instruments
e distinction between binding and non-binding instruments is important. Treaties
(which can alternatively be titled ‘Agreement’, ‘Convention’ or ‘Protocol’), when in
force, impose obligations on their contracting parties that are binding under public
international law. ese legal obligations should be distinguished from the host of
non-binding instruments called ‘Declaration’, ‘Communiqué’, ‘Memorandum of Un-
derstanding’, ‘Action Plan’, ‘Strategy’, ‘Programme’, ‘Initiative’, and the like. Many of
the decisions (Resolutions, Recommendations, etc.) adopted by wildlife treaties’ Con-
ferences of the Parties (COPs – their main decision-making bodies in which all parties
are represented and which meet periodically) are as such non-binding, although they
do have the potential to inuence the interpretation of the binding obligations in the
treaties themselves.
A pertinent example of a non-binding instrument is the Communiqué adopted by
the aforementioned CITES-CMS African lion range state meeting in 2016 (Entebbe
Communiqué), and it is worthwhile to reproduce a selection of the statements it con-
tains. e Communiqué records ‘the main threats (listed in no particular order) for
lions in Africa’ to be:
(1) ‘Unfavourable policies, practices and political factors (in some countries);
(2) Ineective lion population management;
(3) Habitat degradation and reduction of prey base;
(4) Human-lion conict;
(5) Adverse socio-economic factors;
(6) Institutional weakness; and
(7) Increasing trade in lion bones.’
Amongst the recommended measures to counter these threats, the Communi-
qué issues a call on range states to ‘strengthen their legislation on lion conservation
and adopt practices ‘ensuring that agricultural activities and mining operations do
not impede lion conservation.’ Furthermore, and signicantly for present purposes,
the Entebbe Communiqué recognizes ‘the need for transboundary cooperation and
management systems in light of the high number of transboundary lion populations.’
It also emphasizes the notorious ‘lack of resources and capacity,’ which has ‘impeded
the implementation of lion conservation activities on the ground.’ Notably, the Com-
muniqué contains the following statement on the controversial issue of lion trophy
hunting, wherein the 28 range states that attended the meeting:
Highlight the benets that trophy hunting, where it is based on scientically estab-
lished quotas, taking into account the social position, age and sex of an animal, have,
in some countries, contributed to the conservation of lion populations and highlight
the potentially hampering eects that import bans on trophies could have for currently
stable lion populations.’
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
92
Generally, the lion range states call upon ‘CITES, CMS and IUCN to actively
support conservation activities,’ inter alia through the establishment of a ‘mechanism
to develop and implement joint lion conservation plans and strategies, capacity-build-
ing in lion conservation and management,’ and also of a ‘fund for specic emergency
projects for lion conservation.’ In addition, the Communiqué contains several spe-
cic considerations regarding CITES and CMS which will be discussed below. us,
whereas the Entebbe Communiqué is not a legally binding document, it does reect a
consensus amongst 28 range states regarding the threats to lions and the measures to
be taken, which can in turn feed into the application of international wildlife treaties
to lion conservation.
Of particular signicance for present purposes are the two regional Lion Conserva-
tion Strategies that were developed in 2006 for West and Central Africa (IUCN 2006a),
and Eastern and Southern Africa (IUCN 2006b) respectively. ese were prepared by
the IUCN SSC Cat Specialist Group, at the instigation of the 13th CITES COP in 2004,
and with the support of a range of other stakeholders. e Conservation Strategy for the
Lion in West and Central Africa sets out four objectives, together with a range of recom-
mended actions to achieve them: (1) conserve lion habitat in the region; (2) conserve the
lion’s wild prey base; (3) achieve sustainable human-lion coexistence; and (4) reduce the
factors decreasing the viability of lion populations (IUCN 2006a). e overall goal of
the Conservation Strategy for the Lion in Eastern and Southern Africa is to ‘secure, and
where possible, restore sustainable lion populations throughout their present and poten-
tial range’ within the region, ‘recognizing their potential to provide substantial social,
cultural, ecological and economic benets’ (IUCN 2006b). Amongst several objectives
identied to achieve this, the Strategy recommends the development and implementa-
tion of ‘harmonious, comprehensive legal and institutional frameworks that provide for
the expansion of wildlife-integrated land use, lion conservation and associated socio-
economic benets in current and potential lion range’, as well as the alignment of global
legal frameworks such as CITES and CMS with the conservation needs of lions in the
region (IUCN 2006b). At the request of the 11th CMS COP in 2014, the two regional
strategies were reviewed by Bauer et al. (2015). e Entebbe Communiqué adopted by
the 2016 CITES-CMS African lion range state meeting arms that ‘all the objectives of
the Regional Lion Conservation Strategies … remain valid.’ us, even if the strategies
themselves are not legally binding, we note their close ties with the CITES and CMS
legal frameworks in particular, and will revisit their relevance below.
Ramsar Wetlands Convention
In 1971, the Ramsar Convention was adopted in order to ‘stem the progressive en-
croachment on and loss of wetlands’ (Preamble). Wetlands are dened in the Conven-
tion as ‘areas of marsh, fen, peatland or water, whether natural or articial, permanent
or temporary, with water that is static or owing, fresh, brackish or salt, including areas
of marine water the depth of which at low tide does not exceed six metres’ (Article1(1)).
International law and lions (Panthera leo): understanding and improving the contribution... 93
e Ramsar Convention’s central feature is a List of Wetlands of International Impor-
tance, presently comprising over 2,000 sites spread across 169 countries, whereby it
should be noted that many listed wetlands also include dry areas within their bounda-
ries. e Convention’s contracting parties ‘shall formulate and implement their plan-
ning so as to promote the conservation of the wetlands included in the List, and as far
as possible the wise use of wetlands in their territory’ (Article 3(1)). Notably, the latter
half of this obligation applies to all wetlands. ‘Wise use’ of wetlands is understood as
‘the maintenance of their ecological character, achieved through the implementation of
ecosystem approaches, within the context of sustainable development’ (Ramsar COP
Resolution IX.1, 2005). Parties to the Ramsar Convention are also required to ‘promote
the conservation of wetlands … by establishing nature reserves on wetlands, whether
they are included in the List or not, and provide adequately for their wardening’ (Article
4(1)). ey are furthermore expected to cooperate regarding transboundary wetlands
and to ‘coordinate and support present and future policies and regulations concerning
the conservation of wetlands and their ora and fauna’ (Article 5).
Adding sites to the Ramsar List is done principally by the contracting parties them-
selves. Each party must designate at least one site of ‘international importance in terms
of ecology, botany, zoology, limnology or hydrology’ for inclusion in the List (Article 2).
For every candidate site, the domestic authority involved completes a ‘Ramsar Informa-
tion Sheet’ detailing how the site meets the selection criteria, with the Convention Secre-
tariat verifying that it indeed does so. Parties can coordinate the listing of the respective
parts of transboundary wetlands located on their territories, resulting in ‘Transboundary
Ramsar Sites’. Deletions or boundary restrictions of wetlands on the Ramsar List may
be conducted only if an ‘urgent national interest’ of the contracting party involved so
requires, and any associated loss of wetland resources should ‘as far as possible’ be com-
pensated, for instance by creating additional nature reserves (Articles 2(5) and 4(2)). In
order to guide the implementation of the aformentioned legal obligations, a large body
of detailed recommendations regarding the conservation and wise use of wetlands has
been adopted over the years by the Ramsar COP. For instance, the COP has claried that
any harvesting of wildlife (products) from a Ramsar-listed site should be ‘regulated by a
management plan developed in close consultation with the stakeholders,’ and that the
party involved is to ‘ensure that the impact of the harvesting will not threaten or alter the
ecological character of the site’ (Ramsar COP Resolution VII.19, 1999, Annex).
Despite the Convention’s initial emphasis on waterbirds, its broad objectives and
obligations evidently also cover the conservation of other native wild fauna inhabit-
ing wetlands generally, and wetlands on the List in particular. According to one of the
criteria adopted by the COP to guide the selection of wetlands for inclusion in the List
‘a wetland should be considered internationally important if it supports vulnerable,
endangered, or critically endangered species’ (Ramsar COP Resolution VII.11, last
amended by Resolution X.20, 2008). Listed wetlands that are under threat can be in-
cluded in the so-called ‘Montreux Record’, a register of Ramsar sites ‘where changes in
ecological character have occurred, are occurring or are likely to occur’ (Ramsar COP
Recommendation 4.8, 1990).
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94
Ramsar sites of importance to lions
Whereas the Ramsar Convention may not be the rst treaty that comes to mind when
thinking about lion conservation, lions certainly are amongst the beneciaries of wet-
land conservation under the Convention – which currently binds 30 of the 33 lion
range states (Table 2). Whereas lions can survive in very arid regions, home ranges
normally include one or more sources of water. Besides providing water for the lions
to drink, concentrations of prey animals also tend to be above average in riverine or
marshy habitat and around waterholes (Valeix et al. 2010). us, the conservation and
‘wise use’ of such wetlands, even if they are small, is important from a lion conserva-
tion perspective. Notably, the denition of wetlands used under the Convention lacks
a minimum size requirement and includes man-made ones, even ‘farm ponds, stock
ponds, small tanks’ (Ramsar Convention Secretariat 2013), so that pumped water
holes in game reserves are clearly covered, and therefore subject to the ‘wise use’ com-
mitment of Article 3. ere is also no minimum size requirement for listing a site as in-
ternationally important, with the result that even small or temporary sites may qualify
for listing, as may clusters of small sites (Ramsar COP Resolution VII.11, 1999). On
the other end of the scale, some African oodplains and other wetland ecosystems are
so vast that they include many lion home ranges.
Many sites of signicance to lions have been deemed of ‘international importance’
and included in the Ramsar List. Table 3 renders 39 Ramsar-listed sites which are of
actual or potential importance to lions, spread over 19 countries. eir locations are
indicated in Figure 1. ey cover a total surface area of 368,609 km2 (an area larger
than Germany and almost as large as Zimbabwe). Most of these sites (21) are between
1,000 and 10,000 km2. Examples are Parc National des Virunga in the DRC, Eto-
sha Pan in Namibia, Kilombero Valley Floodplain in Tanzania (which overlaps with
the Selous Game Reserve), Kafue Flats and Luangwa Flood Plains in Zambia, and
Mana Pools National Park in Zimbabwe. Eight sites are smaller than 1,000 km2, but
such modest Ramsar sites can still be important for resident lions. Examples include
Uganda’s Murchison Falls-Albert Delta Wetland System (17,293 ha) and Lake George
(15,000 ha), and the Makuleke Wetlands in South Africa (7,757 ha). Ten huge Ramsar
sites cover over 10,000 km2 each, including the Bangweulu Swamps in Zambia, the
Zambezi Delta in Mozambique (> 30,000 km2), and the Okavango Delta System in
Botswana (> 55,000 km2).
In 24 out of 39 cases, the importance of the site for lions, usually alongside other
species, is explicit in the ocial motivation led by the contracting party for listing the
site. (is applies to all sites in Table 3 except Parc National des Virunga; the two 2002
sites in Guinea; Lake Nakuru in Kenya; Estosha Pan in Namibia; Makuleke Wetlands
and St Lucia System in South Africa; Sudd in South Sudan; the two sites in Togo; the
four sites in Uganda; and Kafue Flats in Zambia.) Further, some of the sites in Table3
had, or possibly had, lions when designated but (probably) no longer do. Examples
are La Foret Classée et Réserve Partielle de Faune Comoé-Léraba in Burkina Faso, the
three sites in Guinea, and the two sites in Togo. In such cases, the signicance of the
International law and lions (Panthera leo): understanding and improving the contribution... 95
Table 3. Ramsar sites of importance to lion conservation. Whereas most of these sites currently have
lions, some sites have been included from which lions have disappeared in the recent past. Map codes
indicate the sites’ geographic locations as shown in Figure 1. For more detailed information on each site,
including the reasons for listing and precise location and delimitation, see the Ramsar Sites Information
Service database: http://rsis.ramsar.org/.
Country Ramsar site Size (ha) Since Map
code
Benin Site Ramsar du Complexe W 895,480 2006 1
Zone Humide de la Rivière Pendjari 144,774 2007 2
Botswana Okavango Delta System 5,537,400 1996 3
Burkina Faso
Réserve Totale de Faune d’Arly 134,239 2009 4
La Foret Classée et Réserve Partielle de Faune
Comoé-Léraba 124,500 2009 5
Cameroon Waza Logone Floodplain 600,000 2006 6
Chad Plaines d’Inondation des Bahr Aouk et Salamat 4,922,000 2006 7
Réserve de Faune de Binder-Léré 135,000 2005 8
Democratic Republic of the Congo Parc National des Virunga 800,000 1996 9
Guinea
Niger-Niandan-Milo 1,046,400 2002 10
Sankarani-Fié 1,015,200 2002 11
Gambie-Koulountou 281,400 2005 12
Kenya Lake Nakuru 18,800 1990 13
Mozambique Zambezi Delta 3,171,172 2004 14
Lake Niassa and its Coastal Zone 1,363,700 2011 15
Namibia Etosha Pan 600,000 1995 16
Bwabwata-Okavango Ramsar Site 46,964 2013 17
Niger Parc National du W 220,000 1987 18
South Africa St Lucia System 155,500 1986 19
Makuleke Wetlands 7,757 2007 20
South Sudan Sudd 5,700,000 2006 21
Sudan Dinder National Park 1,084,600 2005 22
Togo Parc National de la Keran 163,400 1995 23
Bassin Versant Oti-Mandouri 425,000 2008 24
Uganda
Lake George 15,000 1988 25
Murchison Falls-Albert Delta Wetland System 17,293 2006 26
Lake Mburo-Nakivali Wetland System 26,834 2006 27
Rwenzori Mountains Ramsar Site 99,500 2008 28
United Republic of Tanzania Malagarasi-Muyovozi Wetlands 3,250,000 2000 29
Kilombero Valley Floodplain 796,735 2002 30
Zambia
Bangweulu Swamps 1,100,000 1991 31
Kafue Flats 600,500 1991 32
Busanga Swamps 200,000 2007 33
Luangwa Flood Plains 250,000 2007 34
Mweru wa Ntipa 490,000 2007 35
Tanganyika 230,000 2007 36
Zambezi Floodplains 900,000 2007 37
Zimbabwe Mana Pools National Park 220,034 2013 38
Victoria Falls National Park 1,750 2013 39
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96
Ramsar designation could, and should, be to safeguard the habitat and prey base of
lions (Sandom et al. 2017) so that recolonization or reintroduction remains a future
option. e same is true of some Ramsar sites in range states where lions are presently
considered extinct. An example is Odzala Kokoua in Congo, which was included in
the Ramsar List in 2012 on the basis of documentation mentioning lions as still pre-
sent within the site. An instance where lions were reintroduced into an area that was
designated a Ramsar site when lions were absent is the St Lucia System in South Africa
(designated in 1986, lions reintroduced in 2013). One site from Table 3 is listed on the
Montreux Record, namely Lake George in Uganda.
Using the Ramsar Convention for lion conservation
Protected areas are crucial to lion conservation. According to Lindsey et al. (2017),
given adequate management, Africa’s protected areas could theoretically support over
80,000 lions – up to four times the total wild lion population remaining in Africa to-
day. Compliance by contracting parties with their legal obligations under the Ramsar
Convention in respect of the sites in Table 3 will thus clearly benet lion conservation.
In practical terms, the Ramsar status of a site and the accompanying international
obligations are likely to be distinct factors inuencing range state authorities, includ-
ing courts, when deciding whether to authorize certain development projects or other
human uses within the site (Gardner et al. 2009). Allowing unsustainable levels of
lion killing or bushmeat poaching would certainly be at odds with parties’ obligations
regarding conservation and ‘wise use’, especially so for sites where lions were part of the
reasons for Ramsar-listing. e inclusion of a site on the Ramsar List thus provides a
layer of protection, in addition to any designations of the area under national legisla-
tion or, indeed, other international instruments.
Added to this is a range of associated benets, such as the development of (more
rigorous) site management plans following listing and the attraction of additional fund-
ing. e latter can be pursued inter alia through the Small Grants Fund established in
1990 to aid developing countries in achieving wetland conservation and the sustain-
able development of local communities depending on wetlands. To illustrate, actions
funded under this scheme have included the development of management plans and
of measures to control wildlife harvesting, for instance patrol vehicles. Gardner et al.
(2009) found that Ramsar-listing for 26 African sites has been instrumental in provid-
ing increased support for protection and management of the sites, scientic studies,
funding opportunities, tourism, and poverty alleviation. Lastly, multinational corpora-
tions, while not legally bound by the terms of the Convention (only states can be con-
tracting parties), can also self-impose commitments towards the conservation of Ram-
sar sites as part of their corporate social responsibility policies. For instance, in 2014
HSBC (one of the world’s largest banking and nancial services holdings) adopted a
policy in which it instructs all its businesses to ‘make appropriate enquiries and not
knowlingly provide nancial services directly supporting projects which threaten the
International law and lions (Panthera leo): understanding and improving the contribution... 97
special characteristics of UNESCO World Heritage Sites or Ramsar Wetlands’ (HSBC
2014). e policy notes that the risks of such irresponsible investments are ‘particularly
high in the forestry, agriculture, mining, energy, property and infrastructure develop-
ment sectors’ (HSBC 2014).
From a lion conservation perspective it seems worthwhile, therefore, to make the
most of the Ramsar Convention as it currently applies to lion habitat, and to promote
the inclusion of further wetlands of importance to lions in the Ramsar List. Examples
of such candidate sites for future Ramsar-listing include Usangu Flats and other wet-
land areas within Ruaha National Park in southern Tanzania, the importance of which
is discussed below.
World Heritage Convention
Broadly similar considerations apply with regard to the other global site-based treaty,
the UNESCO World Heritage Convention (WHC), the purpose of which is to con-
serve both cultural and natural heritage. Many ecologically important areas in Africa
qualify as ‘natural heritage’ as understood in the Convention (Article 2), whereby ‘out-
standing universal value’ from an aesthetic, scientic or conservation point of view is
the common denominator. A selection of these sites has hitherto been included on the
World Heritage List. Unlike the Ramsar Convention, decisions regarding the inclusion
of sites are not made by individual states, but by the World Heritage Committee, the
Convention’s central decision-making body with a rotating membership of 21 states
parties (Article 11). e rst step to be made by a party is to draw up a ‘Tentative List’
of outstanding sites on its territory. From this inventory, it may then proceed to nomi-
nate individual sites formally, whereby it is for the nominating party to demonstrate
the site’s outstanding universal value. e nomination is evaluated by the IUCN in an
advisory capacity, after which the World Heritage Committee takes the nal decision
whether to inscribe the site on the World Heritage List. Whereas most sites are within
a single country, the List also includes a number of transboundary sites.
Each party ‘will do all it can’ to fulll its ‘duty of ensuring the identication,
protection, conservation, presentation and transmission to future generations’ of the
natural heritage on its territory, ‘to the utmost of its own resources’ and, where ap-
propriate, ‘with any international assistance and co-operation’ (Article 4). It is recalled
in this regard that ‘natural heritage’ includes, but is not limited to, sites on the World
Heritage List. Furthermore, to warrant that ‘eective and active measures’ are taken
for the protection of the sites involved, the WHC requires that each contracting party
‘shall endeavor, in so far as possible, and as appropriate for each country,’ to ‘take the
appropriate legal, scientic, technical, administrative and nancial measures neces-
sary for the identication, protection, conservation, presentation and rehabilitation
of this heritage,’ and to ‘integrate the protection of that heritage into comprehensive
planning programmes’ (Article 5). e Operational Guidelines of the WHC instruct
parties to provide for a buer zone when this is necessary for a site’s proper conser-
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
98
vation (World Heritage Committee 2016). A World Heritage Fund (Article 15) is
administered by the Committee to provide targeted assistance for the conservation of
specic sites. e Committee also administers the List of World Heritage in Danger
– the WHC equivalent of the Ramsar Conventions Montreux Record – which ags
sites that are ‘threatened by serious and specic dangers’ (Article 11(4)). Based on
a broad mandate to oversee the implementation of the WHC, the World Heritage
Committee regularly adopts decisions urging particular parties to adopt particular
site-specic measures. As a last resort, the Committee may delete a site from the
World Heritage List altogether.
World Heritage sites of importance to lions
Lions in various parts of Africa prot from the WHC, in a manner broadly similar to the
Ramsar Convention. All of the 33 lion range states except Somalia are currently amongst
the 193 contracting parties to the WHC (Table 2). Table 4 portrays 18 sites, in 15 range
states, which are included in the World Heritage List and which are of actual or poten-
tial importance to lion conservation. eir locations are indicated in Figure 1. For many
of these sites, lions are expressly mentioned in the listing justication. In the aggregate,
the 18 sites cover a surface area of 174,630 km2 (209,453 km2 when including buer
zones). As with Ramsar, most sites (8 out of 18) are between 1,000 and 10,000 km2.
ese include Virunga and Garamba National Parks in the DRC, Niokolo-Koba Na-
tional Park in Senegal, Mana Pools/Sapi/Chewore in Zimbabwe, and the Ngorongoro
Conservation Area in Tanzania – an area which has one of the highest densities of lions
in the world. Four sites are smaller than 1,000 km2, including the transboundary Mount
Nimba Strict Nature Reserve in Côte d’Ivoire and Guinea, and the Kenya Lake System
in the Great Rift Valley. Six of the listed sites are over 10,000 km2 in size, including
the Okavango Delta in Botswana, Serengeti National Park and Selous Game Reserve
in Tanzania, and the recently designated trilateral W-Arly-Pendjari Complex in Niger,
Benin and Burkina Faso – the latter site hosting the sole remaining lion population of
signicance in West Africa. Lions are (probably) gone from some of the sites listed in Ta-
ble 4, such as Comoé National Park and Mount Nimba, but WHC protection can help
keep options open for future reintroduction or recolonization by preserving lion habitat
and prey. Seven of the sites in Table 4 are presently included in the List of World Herit-
age in Danger, while two further sites were temporarily Danger-listed in the past. It will
be noted that various of the World Heritage sites in Table 4 partially or completely over-
lap with Ramsar sites, for instance Virunga National Park (DRC), the W-Arly-Pendjari
Complex, the Okavango Delta (Botswana) and Mana Pools National Park (Zimbabwe).
As for possible future World Heritage listings, Table 5 renders 26 sites which feature in
the Tentative Lists of 14 range states, the successful nomination of which would appear
benecial to lions. Regarding Asiatic lions, the Gir Wildlife Sanctuary was nominated
by India in the past, but the World Heritage Committee decided in 1992 that the site
did not meet the strict criteria for inclusion in the List.
International law and lions (Panthera leo): understanding and improving the contribution... 99
Table 4. Sites on the World Heritage List of importance to lion conservation. Whereas most of these sites
currently have lions, some sites have been included from which lions have disappeared in the recent past.
Map codes indicate the sites’ geographic locations as shown in Figure 1. For more detailed information on
each site, including the reasons for listing and precise location and delimitation, see http://whc.unesco.
org/en/list/. B.z. = buer zone; In danger = listing on List of World Heritage in Danger.
Country World Heritage site Size (ha) Since In danger Map
code
Botswana Okavango Delta 2,023,590
+ b.z. 2,286,630 2014 - 1
Central African Republic Manovo-Gounda St Floris
National Park 1,740,000 1988 1997-present 2
Côte d’Ivoire Comoé National Park 1,150,000 1983 2003-present 3
Côte d’Ivoire & Guinea Mount Nimba Strict Nature
Reserve 18,000 1981 1992-present 4
Democratic Republic of
the Congo
Virunga National Park 800,000 1979 1994-present 5
Garamba National Park 500,000 1980 1984-1992
1996-present 6
Kenya
Lake Turkana National Parks 161,485 1997 - 7
Mount Kenya National Park/
Natural Forest
202,334
+ b.z. 69,339 1997 - 8
Kenya Lake System in the Great
Rift Valley
32,034
+ b.z. 3,581 2011 - 9
Niger, Benin & Burkina
Faso W-Arly-Pendjari Complex 1,494,831
+ b.z. 1,101,221
1996/
2017 -10
Senegal Niokolo-Koba National Park 913,000 1981 2007-present 11
South Africa iSimangaliso Wetland Park 239,566 1999 -12
Uganda Rwenzori National Park 99,600 1994 1999-2004 13
United Republic of
Tanzania
Ngorongoro Conservation Area 809,440 1979 1984-1989 14
Serengeti National Park 1,476,300 1981 -15
Selous Game Reserve 5,120,000
+ b.z. 21,492 1982 2014-present 16
Zambia & Zimbabwe Mosi-oa-Tunya / Victoria Falls 6,860 1989 -17
Zimbabwe Mana Pools National Park, Sapi
and Chewore Safari Areas 676,600 1984 -18
Using the WHC for lion conservation
In parallel to the discussion above regarding the Ramsar Convention, compliance by lion
range states with their obligations under the WHC appears to render distinct advantages
from a lion conservation perspective. For World Heritage sites with lions these obliga-
tions would include the prevention or mitigation of human-lion conict and of prey
depletion. Designation as World Heritage entails signicant prestige, owing in part to
the strict selection criteria and external designation process. is prestigious status puts
real weight in the scales of governmental decision-making regarding activities potentially
aecting listed sites. Likewise, the possibility of a site being stripped of its World Heritage
designation is a signicant incentive for states to comply with their commitments under
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
100
Table 5. Sites of importance to lion conservation on range states’ tentative World Heritage lists. Whereas
most of these sites currently have lions, some sites have been included from which lions have disappeared
in the recent past. For more detailed information on each site, see http://whc.unesco.org/en/tentativelists/.
Country Site on Tentative List Since
Botswana
Chobe Linyanti System 2010
Makgadikgadi Pans Landscape 2010
Central Kalahari Game Reserve 2010
Cameroon Parc National de Waza 2006
Chad Parc National de Zakouma 2005
Ethiopia Bale Mountains National Park 2008
Ghana Mole National Park 2000
Kenya
Lake Nakuru National Park 1999
Aberdare Mountains 2010
e African Great Rift Valley – Hell’s Gate National Park 2010
e African Great Rift Valley – e Maasai Mara 2010
e Great Rift Valley – e Kenya Lakes System 2010
e Meru Conservation Area 2010
Tsavo Parks and Chyulu Hills Complex 2010
Malawi Nyika National Park 2000
Vwaza Marsh Wildlife Reserve 2011
Mali La Boucle du Baoulé 1999
La Réserve de Biodiversité du Parc du Bang Makana 2016
Namibia
Brandberg National Monument Area 2002
Etosha Pan 2016
Okavango Delta 2016
Niger Zone Giraphe 2006
Nigeria Gashaki-Gumpti National Park 1995
Sudan Dinder National Park 2004
Togo Parc National de la Kéran et la Réserve de Faune Oti-Mandouri 2002
United Republic of Tanzania Eastern Arc Mountains Forests of Tanzania 2006
the Convention. is possibility is a ‘stick’ at the disposal of the World Heritage Com-
mittee that the Ramsar Convention lacks. e Committee is also in a position to require
that measures for a site’s protection and management be in place before it is inscribed on
the List – which again is a signicant advantage over the Ramsar Conventions procedure.
Overall, the WHC adds a substantial layer of legal protection and a range of as-
sociated benets in respect of listed sites. For an accessible overview and discussion of
the benets of the WHC for wildlife conservation generally we refer to Bertzky (2014).
Here, we provide a few examples from the past to illustrate the dierent ways in which
the WHC can serve lion conservation. In 1984, the World Heritage Committee de-
cided to include the Ngorongoro Conservation Area in the List of World Heritage
in Danger, after a lack of management had led to the sites overall deterioration. In
subsequent years, thanks in part to the Committee’s active engagement and technical
cooperation projects, the situation improved and the site was removed again from the
Danger List. More recently, the Tanzanian government reversed its plan to upgrade a
International law and lions (Panthera leo): understanding and improving the contribution... 101
road bisecting the Serengeti National Park into a ‘Serengeti Super Highway’, under
pressure from the World Heritage Committee and, in particular, from two rulings of
the East African Court of Justice in 2014 and 2015. In the latter, the Court determined
that upgrading the road would be contrary to Tanzania’s environmental obligations un-
der the EAC Treaty, while leaning heavily on the Serengeti’s World Heritage status in
reaching this verdict (Reference No. 9 of 2010, 20 June 2014; Appeal No. 3 of 2014,
29 July 2015). A nal illustration concerns the role of multinational corporations.
Whereas these are not bound by the WHC as such, an increasing number of them have
undertaken ‘no-go’ commitments regarding sites on the World Heritage List. Besides
the aforementioned HSBC policy, the International Council of Mining and Metals
and oil companies like Shell, SOCO, Total and Tullow Oil have undertaken not to
explore in or extract from World Heritage sites (http://whc.unesco.org/en/extractive-
industries). at recurrent threats of mineral extraction activities in sites like Kenyas
Lake Turkana and the DRC’s Virunga National Park have to date been kept at bay has
been due in large part to these sites’ World Heritage status.
Evidently, the listing of a site on the World Heritage List or the Danger List does
not as such guarantee conservation success. For example, despite its status as a World
Heritage site since 1981 and its Danger-listing in 2007, Senegal’s Niokolo-Koba Na-
tional Park has experienced calamitous declines in prey populations, and concomitant
declines in lion numbers (Henschel et al. 2014). e IUCN estimates the lion popula-
tion has declined by 92%, from over 200 animals to only 16, between 1993 and 2014
(Bauer et al. 2016). Even so, the situation might have been even worse without the
site’s World Heritage status, and that status would also appear to increase the possibili-
ties for promoting recovery.
On the basis of the foregoing, on the whole it appears sensible to seek out and use
the existing opportunities for making the most of the WHC for lions occurring in
extant World Heritage sites, and to actively work towards the future listing of tentative
and other potential heritage sites of importance to lions. One signicant candidate
site, despite not being tentatively listed yet, is Ruaha National Park in southern Tanza-
nia. is largest National Park in East Africa is the core protected area for the world’s
second largest lion population (Dickman et al. submitted; Riggio et al. 2012), and
has very high levels of anthropogenic lion killing on its borders (Abade et al. 2014).
However, it has long been over-looked in terms of its international importance, despite
being highlighted as a Key Landscape for Conservation (KLC) by the European Com-
mission (2016), and as a priority area in international and national lion action plans
(IUCN 2006b; TAWIRI 2007). World Heritage listing could be a welcome improve-
ment of its global recognition and protected status.
Lions as ‘World Heritage species’
As an epilogue to this section, we draw attention to intermittent calls for the intergov-
ernmental recognition of certain species of outstanding universal value as ‘World Her-
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
102
itage species’ (Wold 2008; Wrangham et al. 2008; Hance 2016). Whereas, conceptual-
ly, a good case can be made that lions – alongside other candidates like elephants, tigers
and great apes – are species of ‘outstanding universal value’ and should be considered
part of the world’s common heritage, the WHC currently only provides a legal basis for
declaring sites, not species, as World Heritage. Providing such a legal basis would require
amendment of the WHC or the conclusion of a separate legal instrument dedicated to
World Heritage Species (Wold 2008; see also Arthur 2014).
Convention on International Trade in Endangered Species (CITES)
With the sole remaining exception of newly independent South Sudan, all lion range
states are currently parties to CITES (Table 2). e purpose of the Convention is to
prevent species from being over-exploited through international trade by requiring its
parties to impose restrictions on the international trade of plants and animals (and the
parts and derivatives thereof) which belong to species, subspecies or populations listed
on one of the CITES appendices. Restrictions are implemented through a system of
permits, and the level of restriction corresponds with the level of danger faced by the
species: Appendix I species are threatened with extinction and are therefore subject to
a ban on international commercial trade (Article III); while trade in Appendix II spe-
cies – which are not yet threatened with extinction, but may become so in the absence
of trade regulation – is essentially permissible, provided that it is not detrimental to
the species’ survival (Article IV). Several types of specimens are exempted from CITES’
usual restrictions, including, under certain (complex) conditions, ‘personal or house-
hold eects’, such as hunting trophies (Article VII(3); Res. Conf. 13.7 (Rev. CoP17)).
Captive-bred animals belonging to Appendix I species are treated as if included in
Appendix II (Article VII(4)). More tailored restrictions can be imposed through an-
notations to a species’ listing, which dene the scope of its inclusion in one of the ap-
pendices (Res. Conf. 11.21 (Rev. CoP17)).
While CITES’ legal text is silent on the use of quotas to limit trade in listed
species, the establishment of, and adherence to, quotas is an eective means of sat-
isfying the Convention’s requirement that trade not be detrimental. Quotas can be
established by the COP through either annotation (for instance, the cheetah’s listing
is accompanied by an annotation which expresses annual export quotas for live speci-
mens and hunting trophies from Botswana, Namibia and Zimbabwe) or resolution
(for instance, Res. Conf. 10.14 (Rev. CoP16) recommends quotas for the harvest of
leopards for export from 12 range states). More commonly, however, parties estab-
lish quotas unilaterally at the national level. Parties which fail to comply with their
CITES commitments risk being penalized with trade suspensions (Res. Conf. 14.3),
and, as also tends to be the case with other conservation treaties, parties to CITES
are allowed to adopt domestic measures that are stricter than those required by the
Convention (Article XIV(1)).
International law and lions (Panthera leo): understanding and improving the contribution... 103
CITES, lion hunting trophies, and trade in lion bones and body parts
Given the international movement of hunting trophies and the increasing demand for
lion bone and body parts, CITES clearly has a key role to play in protecting lions against
overexploitation. at said, the divergence between lion population trends in certain
southern African countries and those in the remainder of Africa, combined with the po-
larized nature of the trophy hunting debate (Bauer et al. 2015), have made it challenging
for CITES’ parties to agree on the extent to which trade should be permitted under the
Convention. Since 1977, the Asiatic lion has been listed on Appendix I and the African
lion populations on Appendix II. In addition, three range states (Guinea, Guinea-Bissau
and Somalia) are currently subject to trade suspensions targeting all commercial trade
in CITES-listed species – including lions (http://cites.org/eng/resources/ref/suspend.
php). A growing number of parties, including Australia, the European Union and the
United States, are imposing stricter domestic measures in respect of lions, ranging from
more onerous import requirements than are prescribed by CITES to complete prohibi-
tions on the import of hunting trophies from wild and/or captive-bred animals (CoP17
Prop.4; US Fish and Wildlife Service 2015; see also Macdonald 2016). Declared lion
item exports for the period 2005–2014 numbered 29,214 items, of which 11,164 were
wild sourced (although the denition of wild-sourced is ill-dened, creating some un-
certainty); roughly two-thirds of these items were exported from South Africa – which
has an active captive lion breeding industry (Williams et al. 2015) – with other ex-
porters including Botswana, Ethiopia, Mozambique, Namibia, Tanzania, Zambia, and
Zimbabwe (CoP17 Prop.4). Of these states, only three (Ethiopia, Mozambique and
Zambia) appear to have notied the CITES Secretariat that they use national quotas as
a means of ensuring the sustainability of lion exports (Table 6).
Proposals to up-list the African lion to Appendix I were submitted by Kenya in
2004 (CoP13 Prop.6) and by nine countries from West and Central Africa – all of
which are either currently part of, or have historically belonged to, the lions range – in
2016 (CoP17 Prop.4). In the Entebbe Communiqué, which preceded the 17th CITES
COP in the same year, range states highlighted the importance of considering the lat-
ter proposal against the relevant CITES listing criteria. ey further recognized that:
‘Lion Range States have dierent views on the inclusion of all African popula-
tions of Panthera leo in Appendix I, with some arguing that the populations in West
and Central Africa are fragmented and highly threatened; and others arguing that the
species does not meet the listing criteria and is threatened by factors other than those
CITES can address.’
Following the subsequent negotiations during the 17th CITES COP, the African
lion was ultimately retained on Appendix II. A new annotation was, however, added to
the Appendix II listing, which sets a zero annual export quota for ‘specimens of bones,
bone pieces, claws, skeletons, skulls and teeth removed from the wild and traded for
commercial purposes’, but allows the trade of specimens of bones etc. derived from
South Africa’s captive breeding operations, provided that national export quotas are es-
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
104
Table 6. Unilaterally-set quotas for the export of Panthera leo specimens. Data from http://www.cites.
org/eng/resources/quotas/index.php.
Range State Year Quantity Type of specimen
Ethiopia
2017 10
trophies
2016 10
2015 10
2014 10
2013 5
2012 10
2011 10
2009 20
2008 20
2007 20
80 skins
2006 20 trophies
80 skins
2005 20 trophies
80 skins (conscated)
2004 20 trophies
80 skins
2003 12 trophies
2002 30 trophies
2001 15 live & trophies
2000 10 live & trophies
Mozambique
2017 54
trophies, wild taken2016 54
2015 60
2014 53
wild taken 2013 50
2012 50
Zambia 2017 24 wild taken
2016 24
tablished and communicated to the CITES Secretariat. South Africa has set an export
quota at 800 lion skeletons (Department of Environmental Aairs 2017). e concern
remains that allowing any trade of lion parts is potentially problematic from an en-
forcement point of view and has the potential to stimulate demand, and thus poaching
(Williams et al. 2015). In this regard, the COP retains the discretion to amend this
annotation in the future so as to provide for a more uniform treatment of lion parts
regardless of their origin, or to include further conditions in respect of permissible
trade. It could, for instance, be required that the proceeds of trade be used for lion con-
servation and development initiatives beneting rural communities in lion range, thus
assisting in the mitigation of human-lion conict. A precedent for the latter approach
was set by the annotations restricting trade in elephant ivory.
International law and lions (Panthera leo): understanding and improving the contribution... 105
In addition to its inclusion of a new annotation on the international trade of lion
parts, the 17th CITES COP adopted a series of decisions on the African lion (discussed
below), as well as a resolution on trade in hunting trophies (Res. Conf. 17.9), which
seeks to strike a balance between recognizing the potential benets of trophy hunting
and preventing this practice from occurring at unsustainable levels. In the resolution,
the COP recognizes that ‘well-managed and sustainable trophy hunting is consistent
with and contributes to species conservation, as it provides both livelihood opportu-
nities for rural communities and incentives for habitat conservation, and generates
benets which can be invested for conservation purposes.’ At the same time, the COP
agrees that (even when treated as a personal or household eect) the export of hunting
trophies should generally be conditional upon the issuance of an export permit, and
thus the making of a non-detriment nding. e resolution further provides guid-
ance on the sustainable management of trophy hunting, and recommends, inter alia,
that parties ‘consider the contribution of hunting to a species’ conservation and socio-
economic benets, and its role in providing incentives for people to conserve wildlife,
when considering stricter domestic measures and making decisions relating to the im-
port of hunting trophies’.
Under the current Appendix II listing, African states are limited in the types of lion
specimens that they may export for commercial purposes, and a party which allows
trade to occur at levels that are detrimental to the species’ survival will be in breach of
its CITES commitments. Were all African lion populations ever to be moved to Ap-
pendix I in the future, the types of trade allowed by the Convention would become
even more constrained. However, barring additional restrictions through annotations
or stricter domestic measures, trade in captive-bred lions could continue for commer-
cial purposes. Moreover, as illustrated by CITES’ approach to cheetahs and leopards –
both of which appear on Appendix I – the continued export of hunting trophies would
also be possible, provided that this is not detrimental to the survival of the population
involved. An alternative approach could be to retain some countries’ lion populations
on Appendix II, while shifting the remainder to Appendix I. e COP has already
allowed such ‘split-listing’ for two other members of Africa’s ‘Big 5’ – the African ele-
phant (Loxodonta africana) and the white rhinoceros (Ceratotherium simum) – in order
to accommodate the trade of animals from certain well-managed populations of these
species in southern Africa (see e.g. Lewis 2009). e COP has also, however, cautioned
that split-listing should generally be avoided ‘in view of the enforcement problems it
creates’ (Res. Conf. 9.24 (Rev. CoP17)).
e CITES Animals Committee is tasked with conducting ‘periodic reviews’ of
the species appearing in the Convention’s appendices, with the purpose of advising the
COP on whether particular species are appropriately listed, based on current biological
and trade information in light of the applicable listing criteria (Res. Conf. 14.8 (Rev.
CoP17)). Panthera leowas included in this process in 2011 and, in 2014, a draft review
(suggesting that the African lion’s Appendix II listing remained appropriate (AC27 Doc.
24.3.3)) was presented to the Committee, which considered it necessary to incorpo-
rate information from the lion’s 2015 IUCN Red List Assessment before nalizing the
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
106
document. e review had not been nalized by the 17th COP in 2016, at which stage
the need for its completion fell away as a result of the COP making a decision on the
lion’s proposed up-listing (CoP17 Doc. 82.2). Notably, the CITES COP’s decision not
to uplist the lion was inuenced by the fact that international trade is not the primary
threat faced by the species and that what is needed are consequently not trade bans but
cooperative measures between range states (UNEP/CMS/COP12/Doc.24.3.1.3).
CITES, enforcement issues, and the broader lion conservation agenda
As is highlighted by the COP’s concerns regarding split-listing, CITES’ trade con-
trols clearly cannot be eective unless implemented and enforced (Wandesforde-Smith
2016; Zhou et al. 2016). is is true regardless of the appendix on which a species/
population nds itself. Indeed, in 2002 the CITES COP recognized that, despite the
Appendix I listing of all Asian big cat species (including the Asiatic lion), illegal trade
in these species had escalated and continued to threaten their survival. e COP there-
fore called for a variety of legislative and enforcement measures to address this situation
(Res. Conf. 12.5 (Rev. CoP17)). For Africa’s populations of Panthera leo, it is worry-
ing that 23 of the range states that are parties to the Convention have been assessed
as having inadequate legislation for the eective implementation of CITES (Table 7;
see also Watts 2016). Improvements are clearly desirable in this regard, as are measures
to enhance the capacity of African states to implement and enforce those laws that do
exist (Wandesforde-Smith 2016).
e COP17 decisions on the African lion (Decisions 17.241–245) make no explicit
mention of strengthening national CITES-implementation legislation, but call for a
wide array of measures to improve the conservation and management of this ‘iconic
species’, many of which are clearly responses to the Entebbe Communiqué. Notably,
these CITES COP decisions have also been endorsed by the CMS Standing Committee
and will be presented to the CMS COP for adoption in October 2017 (UNEP/CMS/
COP12/Doc.24.3.1.3). e decisions direct the CITES Secretariat, subject to external
funding and in collaboration with African lion range states, the CMS and the IUCN,
to, inter alia, ‘investigate possible mechanisms to develop and support the implementa-
tion of joint lion conservation plans and strategies, taking into consideration existing
lion conservation plans and strategies’ (the IUCN’s 2006 regional Lion Conservation
Strategies clearly being signicant in this regard); and to take a variety of measures con-
cerning capacity building for joint conservation plans, further international coopera-
tion, ecological and trade research, information-sharing, and education.
Further, the abovementioned decisions direct the CITES Standing Committee
to establish a Task Force on African lions, and to consider establishing a trust fund
to attract funding for both the work of the Task Force and the implementation of
conservation and management plans and strategies for the African lion. Two initia-
tives which seek to defeat wildlife crime in Africa, and whose participation in, or
collaboration with, the African Lion Task Force thus appears to be appropriate, are
International law and lions (Panthera leo): understanding and improving the contribution... 107
Table 7. Status of CITES implementation legislation. Data from http://www.cites.org/eng/legislation,
last updated 01/09/2016.
Category Range state(s)
Category 1
Believed generally to meet all requirements for eective
CITES-implementation
Cameroon, Democratic Republic of Congo,
Ethiopia, Namibia, Nigeria, Senegal,
South Africa, Zimbabwe
Category 2
Believed generally to meet some requirements for
eective CITES-implementation
Benin, Botswana, Burkina Faso, Chad, Guinea,
India, Kenya, Malawi, Mali, Mozambique, Sudan,
Togo, United Republic of Tanzania, Zambia
Category 3
Believed generally not to meet any requirements for
eective CITES-implementation
Angola, Central African Republic, Côte d’Ivoire,
Ghana, Guinea-Bissau, Niger, Rwanda, Somalia,
Swaziland, Uganda
Non-party South Sudan
the Lusaka Agreement Task Force (established by the 1994 Lusaka Agreement) and
the Horn of Africa Wildlife Enforcement Network. Between them, these initiatives
presently cover seven lion range states: Ethiopia, Kenya, South Sudan, Sudan, Tan-
zania, Uganda and Zambia. A nal point concerning enforcement is that the 17th
CITES COP also adopted a resolution on demand reduction strategies as a means
of combatting illegal wildlife trade (Res. Conf. 17.4), prompting some delegates to
question whether it is possible to simultaneously reduce demand for illegal products
and promote the consumption of legal ones, as the resolution on trophy hunting ap-
pears to do (IISD 2016).
Despite its imperfect implementation record and the challenges it faces in balanc-
ing calls for preservation with those for sustainable use (Wandesforde-Smith 2016),
CITES has a demonstrated potential to make a tangible dierence to the conservation
of species threatened by trade. For instance, the conservation status of jaguars (Pan-
thera onca) and other South American felids notably improved after the CITES ban
on trade in their pelts took eect in 1975 (Di Marco et al. 2014). Regarding lions, the
least that can be said is that the relevance of CITES to the conservation and sustainable
use of the species is likely to stay on the increase for some time to come. However, due
to the Convention’s narrow focus on trade, and trade not being amongst the primary
concerns for lion conservation, CITES provides a necessary but not a sucient inter-
national framework for lion conservation.
Convention on Migratory Species (CMS)
e CMS broadly addresses the conservation of migratory species, and like CITES
also lists species in appendices. e Convention supports the conservation and man-
agement of migratory species by requiring that parties take specied conservation
measures in respect of species in CMS Appendix I; by promoting the development of
targeted ancillary instruments, for CMS Appendix II species in particular; and by pro-
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
108
viding a variety of less formal mechanisms for targeting conservation activity towards
particular groups of species or addressing particular cross-cutting threats.
e Convention denes ‘migratory species’ to mean ‘the entire population or any
geographically separate part of the population of any species or lower taxon of wild
animals, a signicant portion of whose members cyclically and predictably cross one
or more national jurisdictional boundaries’ (Article I(1)(a)). is denition allows the
Convention to attach dierent legal commitments to dierent populations of the same
species, and only encompasses wild animals, thus failing to regulate parties’ activities
in respect of animals bred in captivity. Further, the CMS COP has taken a remarkably
exible approach in interpreting the denition, having accepted that taxa which peri-
odically traverse (or have historically traversed) national borders are ‘migratory species’,
even if the reason for these movements is simply that their ranges are transboundary
(Trouwborst 2012). e lion is a case in point. Moreover, lions can disperse over large
distances and some of them migrate along with their migratory prey. In both cases they
may traverse international boundaries (UNEP/CMS/COP12/Doc.25.1.3). However,
the Asiatic lion currently lacks such transboundary features. At any rate, the COP has
explicitly recognised that ‘Panthera leo … and all its evolutionarily signicant constitu-
ents, including Panthera leo persica, satisfy the Convention’s denition of “migratory
species”’ (CMS COP Resolution 11.32, 2014).
Listing lions under the CMS
While CMS Appendix I lists ‘endangered’ migratory species (Article III(1)), Appendix
II is dedicated to migratory species which have an unfavourable conservation status
and require international agreements for their conservation and management, as well
as species whose conservation status, though not necessarily unfavourable, would sig-
nicantly benet from an international agreement (Article IV(1)). At a 2010 meeting
of the Convention’s Scientic Council, Congo, being interested in CMS support for
lion reintroduction eorts, raised the possibility of an Appendix II listing (UNEP/
CMS/ScC16REPORT). In 2014, Kenya submitted a proposal to include the Asiatic
lion on Appendix I and all other subspecies on Appendix II, which was subsequently
revised to propose that all populations of Panthera leo be listed on Appendix II (UNEP/
CMS/COP11/Doc.24.1.2/Rev.1). Kenya’s proposal was ultimately withdrawn, but
the COP adopted Resolution 11.32, which inter alia requested consultations between
range states concerning the population status of Panthera leo, and invited range states,
subject to the ndings of such consultations, to work towards an Appendix II listing
proposal to be presented to the 12th CMS COP in October 2017. Subsequently, in the
Entebbe Communiqué, range states recognized that the ‘CMS can provide a platform
to exchange best conservation and management practices; support the development,
implementation and monitoring of action plans; promote the standardization of data
collection and assessments; facilitate transboundary cooperation; and assist in the mo-
International law and lions (Panthera leo): understanding and improving the contribution... 109
bilization of resources.’ Many range states additionally indicated that they would be in
favour of an Appendix II listing, although southern African states expressed doubt as to
whether their lion populations should be included therein. In accordance with Resolu-
tion 11.32, COP12 is indeed set to consider a proposal for listing the lion in Appendix
II, which was submitted jointly by Chad, Niger and Togo. e proposal, inter alia, de-
scribes how lions may cross national jurisdictional boundaries as part of their circadian
cycles, life cycles, and annual cycles; and identies countries which share lion popula-
tions that are suspected to cyclically and predictably traverse national boundaries, such
that a signicant portion of Africa’s lion population can be considered ‘migratory’ for
CMS purposes (UNEP/CMS/COP12/Doc.25.1.3).
Support for listing lions on CMS Appendix II has also been expressed in the re-
cent literature (Trouwborst 2015a; Watts 2016), and would certainly t the pattern of
prior CMS practice and recent listing trends. e CMS appendices already include the
large carnivore species cheetah and snow leopard (Panthera uncia) in Appendix I, and
African wild dog and polar bear (Ursus maritimus) in Appendix II. e listing propos-
als that will be considered by CMS COP12 include two further carnivores besides
the lion – leopard and Gobi bear (Ursus arctos isabellinus) – as well as other African
megafauna – chimpanzee (Pan troglodytes), girae (Giraa camelopardalis) and African
wild ass (Equus africanus).
In its most recent guidance on assessing proposals to list species on the Conven-
tion’s appendices, the CMS COP has advised, inter alia, that a taxon assessed as ‘Extinct
in the Wild’, ‘Critically Endangered’, ‘Endangered’, ‘Vulnerable’ or ‘Near reatened’
using the IUCN Red List criteria satises the Convention’s denition of ‘unfavourable
conservation status’ and is thus eligible for consideration for Appendix II listing; and
that a taxon assessed as falling into one of the rst three of these categories is eligible for
consideration for listing in Appendix I (Resolution 11.33, 2014). Given their current
Red List categorisations, the Asiatic lion and the West African lion are thus eligible for
CMS Appendix I listing, while the remainder of Panthera leo is eligible for Appendix II
listing. Red List status is not, however, the only relevant consideration. e COP has
also accepted that listing should only occur if this is expected to result in conservation
benets, and has further highlighted the need to consider listing proposals’ ‘coherence
with existing measures in other multilateral fora’ (Resolution 11.33). It is permissible
for species to be listed simultaneously in both Appendices I and II (Article IV(2)).
Should a species that has only been listed in Appendix II decline to the extent that it
becomes endangered, a subsequent Appendix I listing would of course be a possibil-
ity – though by no means a certainty given the COP’s pragmatic approach to listing.
Indeed, 73% of the taxa listed under the Convention appear only in Appendix II
(http://www.cms.int/en/species).
At any rate, were any populations of Panthera leo to be included in CMS Appendix
I, all states belonging to these populations’ current range would become subject to
certain conservation commitments. Although the Convention does not require that
states in which a species is extinct take measures to facilitate its return, any state to
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
110
which the species is reintroduced or which the species (re)occupies spontaneously will,
at that stage, become subject to the same legal requirements as other range states.
ese include the requirement that states endeavour to take measures to conserve and
restore the species’ habitat and address factors which impede its migration or otherwise
endanger the species (Article III(4)); as well as the requirement that taking of animals
belonging to the species be prohibited (Article III(5)). ‘Taking’ in this context includes
‘taking, hunting, ... capturing, harassing, deliberate killing, or attempting to engage
in any such conduct’ (Article I(1)(i)). On the face of it, the requisite taking prohibi-
tion is extremely far reaching, encompassing everything from trophy hunting, to kill-
ing for damage control, to capture for the purposes of research or translocation. e
Convention does, however, allow for certain exceptions – including for scientic pur-
poses, propagation, traditional subsistence use, or where ‘extraordinary circumstances
so require’ (Article III(5)). ese oer CMS parties a measure of exibility and could
conceivably even be relied upon to justify limited trophy hunting, provided that this
is strictly controlled and does not operate to the species’ disadvantage. at said, the
CMS COP has shown a preference for range states in which sustainable taking is
possible to request exclusions from Appendix I listing, rather than to rely upon the
Convention’s exemptions provision (see e.g. Resolution 10.28 on the Saker falcon,
Falco cherrung). Unsurprisingly, there have thus been instances in which the conserva-
tion benets associated with hunting have been relied upon to argue that Appendix I
listing will not be to a population’s benet. For instance, in its assessment of Kenya’s
proposal to list the African lion on Appendix II, the CMS Scientic Council accepted
that, despite the West African lion’s IUCN categorisation as Critically Endangered, an
Appendix II listing seemed the most appropriate course of action, given stakeholders’
belief that a ban on regulated taking would be ‘harmful to the conservation of this
taxon’ (UNEP/CMS/COP11/Inf.8).
Further arguments against certain species’ Appendix I listing have been based
on the permissibility of trade under CITES. For instance, in 2009, three countries
cheetah populations were excluded from the species’ listing on CMS Appendix I be-
cause quotas for trade in these populations are permitted under CITES (UNEP/CMS/
COP9/REPORT). Including the African lion in CMS Appendix I would not interfere
with South Africa’s trade in parts from captive-bred animals. However, such uplisting
would present diculties for states which permit trophy hunting of wild lions. Indeed,
during the Scientic Council’s 2010 discussion of the possibility of listing the African
lion in one of the CMS appendices, the CITES representative highlighted that a CMS
Appendix I listing would raise similar concerns about CITES-compatibility to those
encountered when listing the cheetah (UNEP/CMS/ScC16REPORT). Eight of the
lion’s range states, including states where trophy hunting is practiced, such as Namibia,
are not currently parties to the CMS (Table 2) and therefore would not incur any legal
obligations from an Appendix I listing unless they were to ratify the Convention. Cau-
tion should therefore be taken to consider the positions of these states when making
listing decisions regarding commercially valuable species so as not to deter them from
International law and lions (Panthera leo): understanding and improving the contribution... 111
becoming parties to the Convention. Notably, Botswana, despite being a non-party,
has expressed its support for the CMS Appendix II listing of the African lion (UNEP/
CMS/COP12/Doc.25.1.3). Insofar as the Asiatic lion is concerned, a CMS Appendix
I listing would in fact complement CITES’ ban on the commercial trade of animals
belonging to this subspecies.
CMS ancillary instruments and lions
While the CMS’s substantive conservation requirements only apply in respect of Ap-
pendix I species, the Convention also promotes the development of ancillary instru-
ments, which prescribe detailed conservation measures in respect of particular spe-
cies or groups of species and provide institutional platforms for coordinating, and
reviewing progress towards achieving, such measures. Parties to the Convention must
endeavour to conclude legally binding ‘AGREEMENTS’ for the conservation and
management of Appendix II species (Article II(3)(c)), giving priority to species with
an unfavourable conservation status (Article IV(3)). CMS parties are further encour-
aged to conclude ‘agreements’ in respect of taxa whose members ‘periodically cross one
or more national jurisdictional boundaries’ (Article IV(4)). e latter ‘agreements’,
which oer considerably greater exibility in terms of scope, content and format, have
thus far taken the form of either treaties or non-binding memoranda of understanding
(MoUs). Institutional structures vary from one instrument to the next, but generally
include a management forum (periodic meetings of the parties/signatories), coordina-
tion support (whether provided by the CMS Secretariat, an independent Secretariat,
or a specic state or non-governmental organization), and some form of scientic/
advisory forum (Lee et al. 2010). However, while the legally binding instruments
have the stability provided by core funding, the MoUs by contrast depend ‘exclusively
on voluntary contributions which could be withdrawn or not materialize at any time’
(Lee et al. 2010).
Were Panthera leo or any of its populations to be listed on Appendix II, it would be
possible and in accordance with the Convention to develop a binding AGREEMENT,
whose membership would be open to all range states, regardless of whether they are
CMS parties (Article V(2)). Such an instrument could potentially also incorporate
other large carnivores with overlapping ranges – the African wild dog being an espe-
cially obvious candidate, given its current Appendix II listing and unfavourable con-
servation status (Trouwborst 2015a). Alternatively – and regardless of whether the lion
is ultimately listed on either of the CMS appendices – Article IV(4) would allow the
development of a treaty or MoU focused either exclusively on lions or more broadly
on the conservation and management of transboundary large carnivore populations
throughout Africa and/or Asia (or portions thereof).
On the one hand, there are distinct advantages to providing such a formal, high-
prole and permanent platform in the form of an ancillary instrument, and doing so
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
112
would be in line with the Convention’s provisions. On the other hand, the develop-
ment and functioning of a new ancillary instrument entails administrative and nan-
cial burdens. As with any international legal instrument, this can be expected to inu-
ence states’ willingness both to initiate the development of, and become parties or (in
the case of an MoU) signatories to, such an instrument. Given the urgent need to direct
resources towards in situ conservation eorts, states are likely to be especially hesitant
to develop a new instrument, with an independent administrative and/or decision-
making structure, if they consider it possible to achieve their objectives under existing
legal and institutional frameworks. Indeed, in the face of resource constraints, the CMS
COP has recognized the need to avoid an unwarranted proliferation of ancillary instru-
ments and has adopted criteria against which to assess proposals for the development
of new instruments (Resolution 11.12, 2014). One such criterion, quite sensibly, is the
absence of superior alternatives – either outside the CMS system or within it.
CMS Concerted Actions and lions
One type of alternative remedy within the CMS system is the establishment, through
resolution, of ‘Concerted Actions’ to improve the conservation status of specied Ap-
pendix I and II species, the implementation of which is monitored by the Conven-
tion’s Scientic Council (Resolution 10.23, 2011). Concerted Actions may operate on
a single- or multi-species basis and the COP has accepted that they may act as either
a precursor or alternative to the conclusion of a dedicated treaty or MoU (Resolu-
tion 11.13, 2014). e Scientic Council has recognized that, if listed on either of
the CMS appendices, the lion would be an appropriate species for Concerted Action
(UNEP/CMS/COP11/Inf.8).
In addition, portions of the lion’s present and historic range are already encom-
passed by two existing, geographically-based, multi-species Concerted Actions: the Sa-
helo-Saharan Megafauna Concerted Action and the Central Eurasian Aridland Mam-
mals Concerted Action. e species on which these Concerted Actions are initially
centred include two species of large carnivores – snow leopard and cheetah – and
the COP’s intention is that they ‘will in due course cover all threatened migratory
large mammals of the temperate and cold deserts, semi-deserts, steppes and associated
mountains’ of the Sahelo-Saharan region and Central Eurasia (Recommendations 9.1
and 9.2, 2008). Importantly for the Asiatic lion, the COP has requested the Scientic
Council and the Secretariat to ‘ensure that all means that can eectively contribute to
an improvement of the conservation status of Asian big cats and to awareness raising
on the threats they face are taken within the framework of the Central Eurasian Ar-
idland Mammals Concerted Action’ (Recommendation 9.3, 2008). Lion populations
not falling within the geographic scope of the existing multi-species Concerted Actions
could theoretically be covered by a Sub-Saharan Megafauna Concerted Action, the es-
tablishment of which has already been identied as a possibility by the CMS Scientic
Council (UNEP/CMS/COP11/Inf.8).
International law and lions (Panthera leo): understanding and improving the contribution... 113
CMS Action Plans, Special Species Initiatives and lions
Species action plans can play a key role in operationalizing Concerted Actions.
However, such plans can also be developed, or existing plans endorsed (the regional
Lion Conservation Strategies being potential candidates), within other contexts within
the CMS regime. So can international working groups to monitor and support their
implementation. A further available mechanism takes the form of ‘Special Species
Initiatives’, the prime example being the Central Asian Mammals Initiative (CAMI).
e CAMI and its associated Programme of Work, the implementation of which is
coordinated by the CMS Secretariat, act as a common strategic framework for action,
drawing together the various CMS instruments and mandates of relevance to the species
involved (Resolution 11.24, 2014). e establishment, in collaboration with the CITES
Secretariat, of a similar initiative for African carnivores will be proposed at this year’s CMS
COP. It is envisaged that this Joint CMS-CITES African Carnivores Initiative will be
used to develop both ‘concrete, coordinated and synergistic conservation programmes’
and ‘policy guidance and recommendations’; and to ‘organize the collaboration with
other conservation initiatives and organizations’ (UNEP/CMS/COP12/Doc.24.3.1.1).
While the CAMI focuses primarily on Concerted Action species, four of the 15 species
it covers are not listed on the CMS Appendices. is suggests that it would be possible
for the Asiatic lion to be incorporated into the Initiative, even without CMS listing.
It similarly suggests the possibility of the anticipated African Carnivores Initiative to
encompass not only listed, but also non-listed species.
Flexibility and limited resource demands are amongst the advantages of Concerted
Actions and Special Species Initiatives, and securing the initial participation of states
may also be easier than with a binding ancillary instrument. Conversely, compared to
an ancillary treaty, it may be harder to maintain states’ commitment and to monitor
implementation over time, due to a lack of core funding, a dedicated institutional
structure and ‘legal teeth’.
As a nal and more general point, whereas it is clear from the above that the CMS
regime oers certain options for directing conservation action towards non-listed spe-
cies, listing the lion on either or both of the Convention’s appendices would raise
the species’ prole and would signicantly increase the likelihood of lions being af-
forded priority within the Conventions busy agenda. Indeed, the CMS Secretariat
has observed that it may not be justiable to dedicate the Convention’s limited re-
sources to supporting the conservation of an unlisted species (UNEP/CMS/COP12/
Doc.24.3.1.3).
Convention on Biological Diversity (CBD)
All 33 lion range states are contracting parties to the CBD, which aims broadly for the
conservation and sustainable use of biological diversity, including at the ecosystem,
species and genetic level. e Convention lacks lists of species requiring special atten-
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
114
tion. Regardless, many of the duties it spells out are of plain relevance to lions. ese
include obligations regarding national biodiversity strategies, plans or programmes
(Article 6), in-situ conservation (Article 8), sustainable use (Article 10) and environ-
mental impact assessment (Article 14). To single out one of these, Article 8 requires
each party, ‘as far as possible and as appropriate’, inter alia to establish a ‘system of pro-
tected areas or areas where special measures need to be taken to conserve biological di-
versity’, ‘[p]romote the protection of ecosystems, natural habitats and the maintenance
of viable populations of species in natural surroundings’, ‘[r]ehabilitate and restore
degraded ecosystems and promote the recovery of threatened species’, and ‘[d]evelop or
maintain necessary legislation and/or other regulatory provisions for the protection of
threatened species and populations’. Whereas the above provisions are just as binding
as other treaty obligations, they are phrased in such a broad and qualied manner that
it is dicult in practice to identify the boundary between compliance and violation.
Parties evidently dispose of an ample margin to determine what, in their individual
circumstances, is ‘possible’ and ‘appropriate’, although this discretion is not limitless.
For instance, allowing a species to go extinct on its territory is clearly hard to reconcile
with a state’s obligations under the CBD.
For present purposes, the CBD is also of signicance as a high-prole forum for
signaling, discussing, and sharing information and experience regarding all manner
of conservation issues; as a catalyst for mainstreaming the consideration of biodiver-
sity into broader policy agendas; and as a source of non-binding but authoritative
guidance as developed and endorsed by the CBD COP. Most of the strategic ‘Aichi
Biodiversity Targets’ adopted by the COP in 2010, for instance, are relevant to lion
conservation, such as the 12th: ‘By 2020 the extinction of known threatened species
has been prevented and their conservation status, particularly of those most in decline,
has been improved and sustained’ (CBD Strategic Plan for Biodiversity 2011–2020).
Also of evident relevance are the 2004 ‘Addis Ababa Principles and Guidelines for the
Sustainable Use of Biodiversity’, according to which it is ‘possible to use biodiversity
components in a manner in which ecological processes, species and genetic variability
remain above thresholds needed for long-term viability,’ while ‘all resource managers
and users have the responsibility to ensure that use does not exceed these capacities
(CBD COP Decision VII/12, 2004).
Given the threat posed by depletion of lions’ prey base, the CBD’s active role in
addressing the unsustainable use of bushmeat is particularly relevant. e Conventions
Liaison Group on Bushmeat has developed specic recommendations to complement
the Addis Ababa Principles and Guidelines in this regard, which have been endorsed
by the CBD COP (CBD COP Decisions XI/25, 2012, and XII/18, 2014), and also
by the CITES COP (Res. Conf. 13.11(Rev. CoP17)). e CBD COP has urged par-
ties to develop and promote methods and systems, and build capacity and community
awareness ‘to determine sustainable wildlife harvest levels at national and other levels,
with a particular view to monitoring and improving sustainable wildlife management
and customary sustainable use,’ and to develop and promote ‘sustainable alternatives
to the unsustainable use of wildlife’ (CBD COP Decision XI/25, 2012). Bushmeat is
International law and lions (Panthera leo): understanding and improving the contribution... 115
furthermore addressed in a volume of the CBD Technical Series (Nasi et al. 2008) and,
pursuant to COP Decision XI/25, a Collaborative Partership on Sustainable Wildlife
Management (CPW) was established, which has developed a sourcebook on bush-
meat. Notably, the CPWs 14 members include both CITES and the CMS, and the
latter’s 2017 COP will consider the adoption of several draft decisions on addressing
the unsustainable use of wild meat (UNEP/CMS/COP12/Doc.24.4.7).
Regional instruments
In addition to the global conventions considered above, here we summarize several
relevant regional agreements, although we stress that this concise treatment does not
necessarily reect a lesser practical importance of these instruments to lions.
African Convention
e 1968 African Convention, administered by the African Union, is in force for 21
lion range states (Table 2). Notably, Botswana, Namibia, South Africa and Zimbabwe
are not amongst its contracting parties. e lion – alongside six other large carni-
vores – is listed as a protected species in the Annex to the Convention. Consequently,
contracting parties are under an obligation to ensure that lions are ‘totally protected’
throughout their territories, which includes prohibiting their hunting, killing and cap-
ture (Article VIII). As lions are subject to the exible ‘Class B’ regime, this prohibi-
tion may be lifted ‘under special authorization’ at the discretion of the ‘competent
authority’. e Convention places restrictions on certain means of capture and killing,
including a prohibition on the use of poisoned baits (Article VII). Trade in lions and
lion trophies must be regulated, and their export, import and transit made subject to
an authorization ‘which shall not be given unless the specimens or trophies have been
obtained legally’ (Article IX). Regarding lion habitat, the Convention requires par-
ties to maintain, expand and/or newly establish ‘conservation areas’ – a term covering
‘strict nature reserves’, ‘national parks’ and ‘special nature reserves’ – so as to ‘ensure
conservation of all species and more particularly of those listed … in the annex’ (Arti-
cle X(1)). Concerning the peripheries of such protected areas, parties ‘shall establish,
where necessary, around the borders of conservation areas, zones within which the
competent authorities shall control activities detrimental to the protected natural re-
sources’ (Article X(2)).
e African Convention appears to have contributed to the increase in protected
areas and improvements in national hunting and wildlife trade legislation in many
lion range states during the years following the Conventions adoption (Bowman et al.
2010). Unfortunately, however, the failure of the Convention’s drafters to establish a
COP or similar institutional framework to oversee and promote implementation and
enforcement has made the 1968 African Convention something of a ‘sleeping treaty
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
116
(Bowman et al. 2010). A substantially revised version of the Convention – including
an institutional framework but lacking a species-specic focus – was negotiated in
2003, but requires a further two ratications to enter into force.
Bern Convention
e Bern Convention, the Council of Europe’s counterpart of the African Convention,
is something of an oddity in the current review. Notwithstanding its primary focus on
European wildlife, as reected in its title, in certain ways the geographic scope of the
Convention extends beyond Europe. Without going into the particulars (see Bowman
et al. 2010), we note here that the Bern Convention has a small number of African
states parties, including two lion range states, Burkina Faso and Senegal. e lion itself
is not listed under the Convention – although leopard, tiger (Panthera tigris) and dhole
(Cuon alpinus) are (see also Trouwborst 2017). Still, it would seem that the general
obligation in Article 2 of the Bern Convention requires Burkina Faso and Senegal to
‘take requisite measures to maintain the population of [lions] at, or adapt it to, a level
which corresponds in particular to ecological, scientic and cultural requirements’ –
i.e., a level at which the population is not threatened with extinction (Bowman et al.
2010; Trouwborst et al. 2017b). Interestingly, in 2005 the Standing Committee (the
Bern Convention’s COP equivalent) called for increased international cooperation re-
garding transboundary populations of large carnivores, including: ‘Lion (Felis leo) and
leopard (Panthera pardus) in the National Park of Niokolo Koba (Senegal) and Mali
(Standing Committee Recommendation No. 115, 2005). Overall, however, the rel-
evance of the Bern Convention to lion conservation appears to have been marginal at
best, and there are no indications for this to radically change in the foreseeable future.
SADC Protocol on Wildlife Conservation and Law Enforcement
e SADC covers the large region from the tip of South Africa to the DRC and Tanzania
in the north. e SADC Protocol on Wildlife Conservation and Law Enforcement is cur-
rently in force for eight lion range states (Table 2), and could in future apply to a further
three range states once they ratify (Angola, DRC, Swaziland). e Protocol is intended to
provide ‘common approaches to the conservation and sustainable use of wildlife resources
and to assist with the eective enforcement of laws governing these resources’ (Article
4(1)), whereby ‘wildlife’ is dened as ‘animal and plant species occurring within natu-
ral ecosystems and habitats’ (Article 1). Some of the Protocol’s specic objectives are to
promote sustainable wildlife use; harmonize relevant legal instruments; assist in national
and regional capacity-building for wildlife conservation, management and law enforce-
ment; facilitate community-based management practices; and to promote conservation of
shared wildlife populations through the establishment of TFCAs (Article 4(2)). To achieve
these objectives, the Protocol lays down a range of obligations, accompanied by an institu-
International law and lions (Panthera leo): understanding and improving the contribution... 117
tional framework. e latter includes a Committee of Ministers, a Committee of Senior
Ocials, a Technical Committee composed of the Directors of countries’ wildlife agen-
cies, and a ‘Wildlife Sector Technical Coordinating Unit’ acting as Secretariat (Article 5).
Whereas the Protocol does not contain species-specic provisions, many obliga-
tions are of signicance from a lion conservation perspective. For instance, each con-
tracting party ‘shall ensure the conservation and sustainable use of wildlife resources
under its jurisdiction’ (Article 3(1)). To that end, parties ‘shall adopt and enforce legal
instruments’ (Article 6(1)) and ‘assess and control activities which may signicantly af-
fect the conservation and sustainable use of wildlife so as to avoid or minimise negative
impacts’ (Article 7(2)). Parties shall take measures to ‘ensure the maintenance of viable
wildlife populations’ and prevent over-exploitation, including by regulating the taking
of wildlife through ‘restrictions on the number, sex, size or age of specimens taken and
the locality and season during which they may be taken’ (Article 7(3)). Regarding trans-
boundary populations, parties shall, as appropriate, ‘establish programmes and enter
into agreements to promote the co-operative management of shared wildlife resources
and wildlife habitats across international borders’ (Article 7(5), and generally ‘promote
the development of transfrontier conservation and management programmes’ (Arti-
cle 7(9)). Likewise, parties are to ‘endeavour to harmonise national legal instruments
governing the conservation and sustainable use of wildlife resources’ (Article 6). A par-
ticularly important instrument to further the coordination and harmonization of the
management of transboundary wildllfe populations and ecosystems is the establish-
ment of TFCAs (discussed below). Lastly, we highlight the development of thematic
international strategies developed within the framework of the SADC Protocol, such
as the SADC Law Enforcement and Anti-Poaching Strategy 2016–2021.
In sum, the relevance of the Protocol to ensuring conservation and sustainable
use of lions in the SADC region is evident. We do draw attention to the diculties
involved in implementing the various objectives and obligations in the Protocol. For
instance, the transboundary harmonization of legislation can be quite a challenge, as
illustrated by the analysis conducted by Selier et al. (2016) regarding the management
of a trilateral elephant population in the SADC region.
Treaties establishing Transfrontier Conservation Areas
Some particularly signicant treaties from a lion conservation viewpoint have a mod-
est geographic scope. ese are the bilateral or trilateral treaties establishing TFCAs,
although one exceptional treaty involves ve parties. Four treaty-based TFCAs of im-
portance to lions are:
Kgalagadi (Botswana, South Africa)
Great Limpopo (Mozambique, South Africa, Zimbabwe)
Kavango Zambezi (Angola, Botswana, Namibia, Zambia, Zimbabwe)
Malawi-Zambia (Malawi, Zambia)
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
118
Another four TFCAs of actual or potential importance to lion conservation are as
yet based only on MoUs:
Lubombo (Mozambique, South Africa, Swaziland)
Iona Skeleton Coast (Angola, Namibia)
Greater Mapungubwe (Botswana, South Africa, Zimbabwe)
Chimanimani (Mozambique, Zimbabwe)
TFCAs which are still to be formalized include:
Liuwa Plains-Mussuma (Angola, Zambia),
Lower Zambezi-Mana Pools (Zambia, Zimbabwe)
ZiMoZa (Mozambique, Zambia, Zimbabwe)
Kagera (Rwanda, Tanzania, Uganda)
Niassa-Selous (Mozambique, Tanzania)
Mnazi Bay-Quirimbas (Mozambique, Tanzania)
(For the latest developments regarding each TFCAs, see http://www.peaceparks.org.)
For illustrative purposes, we discuss one TFCA treaty, selecting the most spectacu-
lar one. In 2011, the presidents of Angola, Botswana, Namibia, Zambia and Zimba-
bwe concluded the Treaty on the Establishment of the Kavango Zambezi Transfrontier
Conservation Area (KAZA TFCA), which entered into force a year later. e resulting
TFCA encompasses and unites a huge array of pre-existing protected areas and multiple
resource use areas in the ve countries, many of which are important lion areas, and cur-
rently covers approximately 520,000 km2 – roughly the size of France. While duly rec-
ognizing its ties with the SADC (Article 9), the Treaty formally established the KAZA
TFCA as an autonomous ‘international organisation’ with legal personality (Article 3),
and headquarters in Kasane (Article 2). e Treaty set up various institutions charged
with administering and further developing the KAZA TFCA, including a Ministerial
Committee, Committee of Senior Ocials, Joint Management Committee, Secretariat
and National Committees (Articles 10-23; see also http://www.kavangozambezi.org).
e KAZA TFCA aims to ‘maintain and manage’ the shared natural resources and
biodiversity of the area to ‘support healthy and viable populations of wildlife species’,
and to develop a ‘complementary network of Protected Areas within the KAZA TFCA
linked through corridors to safeguard the welfare and continued existence of migratory
wildlife species’ (Article 6(1)). Other objectives of relevance to lions are to transform
the TFCA into a ‘premier tourist destination in Africa’; to enhance the sustainable use
of natural resources to improve human livelihoods and reduce poverty; to ‘promote
and facilitate the harmonisation of relevant legislation, policies and approaches’; and to
‘ensure compliance with international protocols and conventions related to the protec-
tion and Sustainable Use of species and ecosystems’ (Article 6(1)).
e general principles that the ve states are expected to uphold in their pursuit
of these objectives include the recognition that the right to utilize natural resources
International law and lions (Panthera leo): understanding and improving the contribution... 119
‘carries with it the obligation to do so in a responsible manner so as to ensure eective
Conservation and management for posterity;’ to ensure that wildlife use is sustainable;
to rehabilitate declining populations; and generally to take ‘knowledge based deci-
sions derived from interdisciplinary research and traditional knowledge and to exercise
precaution when there is insucient information’ (Article 5). e ve partner states
are under obligations to ‘ensure the protection and management of those parts of the
Kavango Zambezi ecosystem falling directly under their jurisdiction;’ to cooperate in
developing common approaches to inter alia wildlife management and tourism; and to
ensure proper stakeholder engagement at national and local levels (Article 8).
In sum, investing in the implementation of existing TFCA treaties and the adop-
tion of treaties for further areas can evidently be benecial for lion conservation and
sustainable use. Consolidating the Niassa-Selous TFCA would seem particularly im-
portant, as this area hosts the largest lion population, estimated at over 5,000 lions
(Dickman et al. submitted).
Discussion and recommendations
e above review reveals a signicant body of international wildlife law of relevance
to the conservation and sustainable use of lions. Moreover, it reveals a signicant
potential for enhancing the contribution of wildlife treaties in this regard. e time
is right to invest in such improvements, and our analysis renders several general
and treaty-specic recommendations for doing so. Some of the most signicant are
provided below.
It is appropriate to place our ndings in perspective by noting that no number or
combination of relevant treaties can by themselves secure the conservation of lions into
the long-term future. International wildlife law provides one set of tools in a much
larger toolkit comprising a range of other approaches, mechanisms and disciplines,
many of which are likely to be needed.
Implementation and participation
It seems safe to assume that the future of lions would be much more secure if all range
states fully lived up to the international obligations identied in the above analysis.
However, the implementation of these obligations is aected by pervasive compliance
deciencies due to problems of capacity, governance and enforcement in many range
states (Dickman et al. 2015; Dickman et al. submitted). All eorts aimed at decreasing
these deciencies and improving compliance are thus to be encouraged.
Figure 3 shows a summary of Dickman et al’s (submitted) index of infrastructural
fragility for the 33 lion range states. In brief, this index is based on a set of socio-
political, habitat and conservation variables that are likely to inuence the success of
conservation measures to secure lions within each range state. us, the geopolitical
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
120
score is the sum of standardized (relative to the average of the sample states), national-
level data on factors including: the level of political corruption, stability and regula-
tory quality (governance metrics), measures of economic development (GDP and the
Human Development Index), human population growth and density (factors that put
pressure on available lion habitat) and the percentage of land designated as protected
area (conservation).
Depending on the particular circumstances and the treaty obligation(s) involved,
there is a time and a place for top-down as well as bottom-up approaches, for coercive
as well as exible approaches, and for all manner of combinations of these (Treves et
al. 2017; Chapron et al. 2017; Redpath et al. 2017). It is important to note in this
regard that the participation of local and indigenous communities, poverty alleviation,
awareness raising and education have become key features in the implementation of
all the major conservation treaties, as expressed in COP decisions, strategies, funding
allocations, and guidance documents (see, e.g., Ramsar Convention Secretariat 2010;
CBD Secretariat 2011; UNESCO et al. 2012).
Another generic issue is that of participation gaps at the intergovernmental level.
e utility of some treaties to lions could be improved through the accession of range
states that are still missing as contracting parties, such as Botswana, Namibia and Zam-
bia in the case of the CMS. Further participation gaps are indicated in Table 2.
Figure 3. e map shows the geopolitical values of lion range states, where higher values represent greater
fragility in the infrastructure of the state (based on Dickman et al., submitted).
International law and lions (Panthera leo): understanding and improving the contribution... 121
Site-based treaties
For sites of importance to lions that are listed under the Ramsar and/or World Heritage
Convention, it is clearly worthwhile to take advantage of that international status in order
to improve site management and avert harmful human impacts, as appropriate. e oppor-
tunities to do so are wide-ranging, varying from the Conventions’ funding schemes to liti-
gation, and will generally be greatest for sites where lions were part of the ocial motivation
for listing. e possibility of listing on the Montreux Record and/or List of World Heritage
in Danger can also provide useful leverage. For listed sites from which lions have disap-
peared we recommend not losing sight of lions in site management but rather enabling and
working towards their future return, in particular by conserving their habitat and prey base.
Likewise, there is clear merit in working towards the listing of additional sites of
importance to lions under either Convention. For the World Heritage Convention,
range states’ tentative lists would be the natural starting point in this connection (see
Table 5 for candidates), although some important candidate sites are not yet on these
lists – Tanzania’s Ruaha National Park being a case in point. Signicantly, the listing of
transboundary sites is eligible under both the Ramsar and the World Heritage Conven-
tions. e proper conservation and management of transboundary sites for lions can ev-
idently also benet substantially from their designation as a TFCA through a dedicated
treaty. Such a TFCA agreement can also assist in implementing applicable international
obligations under other instruments for the sites in question. e consolidation of the
Niassa-Selous TFCA is of particular importance from a lion conservation perspective.
Generally, the more international designations a site has, the better its chances of
survival and appropriate management. Ramsar designation is easier to achieve than
World Heritage listing, although once achieved the latter status is of a higher legal
caliber (and is available for a broader range of habitats). Ramsar designation can also
be an intermediate step towards ultimate World Heritage listing.
Both for existing and potential future sites with an international designation, it is
essential to address the unsustainable killing of lions and their prey not only within
but also around the borders of those sites, and to avoid simply relocating human-lion
conict to the sites’ peripheries.
Species-based treaties
CITES provides a necessary framework for trade-related threats to lions and there re-
mains scope to strengthen the Convention’s restrictions, as necessary, either by uplist-
ing African lion populations to Appendix I or adding further annotations to the current
Appendix II listing. If established, the joint CITES-CMS African Carnivores Initiative
will provide an opportunity to address problems aecting implementation. However,
CITES does not provide sucient mechanisms for addressing threats other than trade.
Regarding the CMS, our review indicates that there is denite scope and need for
reinforcement and coordination of actions to further lion conservation and sustainable
Arie Trouwborst et al. / Nature Conservation 21: 83–128 (2017)
122
use across the species’ range. All the other treaties we reviewed appear to be of actual or
potential use in this regard, and sometimes contribute crucial pieces of the puzzle. Yet,
all of them are subject to limitations. e Ramsar Convention is limited to wetlands;
the WHC is limited to sites of outstanding universal value; CITES is limited to inter-
national trade; the CBD is very general and lacks a species-specic focus; the African
Convention is institutionally dormant and several important range states are not par-
ties; the Bern Convention is of marginal signicance; the SADC Protocol has a limited
geographic scope and lacks a species-specic focus; and the various TFCA treaties have
geographically limited and fragmented scopes, and remain conceptual in some of the
most signicant habitats for lions. Given the fragmented collection of treaties which
currently apply to lions and the absence of adequate international instruments and/or
institutions for lion conservation in at least portions of the species’ range, an important
role appears, in principle, to be reserved for the CMS, both in terms of coordination
and gap-lling. Listing lions under the Convention would be a logical step in this re-
gard, and our analysis provides strong support for doing so.
e species’ currently proposed listing on Appendix II would both signal the need
to develop more elaborate species-specic frameworks for lion conservation and sustain-
able use and increase the avenues available for achieving this. Should CMS COP12 de-
cide to list the lion or any of its populations on the CMS Appendices, it would further
seem sensible for the COP to designate lions for Concerted Action – whether this be as
a precursor to the eventual development of an ancillary instrument or as an alternative
thereto. Concerted Action for the Asiatic lion could, in principle, be implemented by
including this subspecies in CAMI. For Africa, the proposed CMS-CITES African Car-
nivores Initiative has the potential to enhance coordination and collaboration amongst
existing conservation initiatives and instruments throughout the African lion’s range
and could play an especially pronounced role in subregions which lack alternative trea-
ty mechanisms to support transboundary cooperation and national implementation.
Indeed, the establishment of coordination and support mechanisms under the CMS
should, in principle, assist range states to implement legal commitments which they
have long held under other international instruments (such as the African Convention),
regardless of whether or not they at some stage decide to undertake new legal commit-
ments through CMS Appendix I listing or the development of an ancillary treaty.
Concluding observations
With their long-term, legally binding commitments on a transboundary scale; their
high proles; their platforms for cooperation and coordination; and various support
mechanisms, international treaties have a distinct contribution to make to lion conser-
vation. e above review makes clear what can and cannot be expected of international
wildlife law in this regard. Importantly, our review shows that there is still much to be
gained, partly by advancing the eective implementation of the currently applicable
law in the diverse and often challenging domestic contexts of the various lion range
states, and partly by enhancing the legal framework itself.
International law and lions (Panthera leo): understanding and improving the contribution... 123
At the intergovernmental level, listing lions under the CMS can be expected to
render particular advantages in terms of the coordination and facilitation of lion con-
servation action across the species’ range. Other recommendations owing from our
analysis include making optimal use of the World Heritage and Ramsar Conventions,
CITES and TFCA treaties for lion conservation. Overall, in order to maximize range
states’ compliance with their international commitments concerning lions, the devel-
opment and implementation of participatory conservation strategies adjusted to na-
tional and local circumstances appears crucial.
Acknowledgements
AT acknowledges the support of the Netherlands Organization for Scientic Research,
as part of the project Ius Carnivoris (project no.: 452-13-014). AJD undertook this
work while a Kaplan Senior Research Fellow and EAM was Rivington Winant Re-
search Fellow in Conservation. e authors thank Clara Nobbe for providing informa-
tion on the CMS Secretariat’s recent lion-related activities, and thank the reviewers for
their helpful comments.
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... Hitherto, such research concerning terrestrial megafauna has predominantly addressed large carnivores and some of the megaherbivores. Species-specific international law analyses regarding large carnivores have focused inter alia on lions Panthera leo (Watts 2016;Trouwborst et al. 2017b;Bauer et al. 2018;Hodgetts et al. 2018), cheetahs Acinonyx jubatus (Nowell and Rosen 2018), leopards Panthera pardus (Trouwborst et al. in press), and ocelots Leopardus pardalis (Kimberley 2017), in addition to an extensive legal literature on gray wolves Canis lupus and other large carnivores in Europe (see, e.g., Epstein 2013;Linnell et al. 2017;Trouwborst 2018). The international law literature on large herbivores includes research regarding hippopotamus Hippopotamus amphibius (Snyder 2015), rhinoceroses (Leader-Williams et al. 2005;Ayling 2013;Coetzee and Couzens 2017;Janssens and Trouwborst 2018), and especially elephants (Glennon 1990;Kidd and Cowling 2003;Couzens 2014;Nollkaemper 2014;Selier et al. 2016;Persaud 2017;Wandesforde-Smith 2016;Jung 2017), with most studies predominantly addressing international trade regulation. ...
... The text and context of the various While identifying and illustrating the relevance of various legal instruments for large herbivore conservation, no detailed descriptions of the actual legal obligations of contracting parties, institutional structures, general functioning and overall conservation benefits of the various treaty regimes are given below, to avoid undue repetition. A possible starting point when looking for such detail is Trouwborst (2015), discussing most of the instruments mentioned below with respect to large carnivores; other suggested sources are Bowman et al. (2010Bowman et al. ( , 2016, Gillespie (2011), Trouwborst et al. (2017b, and Janssens and Trouwborst (2018). ...
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... Large carnivores play vital ecological roles as apex predators (Estes et al. 2011, Ripple et al. 2014; thus, their extirpation may have cascading effects on ecosystem structure and functioning. Due to their considerable humanwildlife conflict potential and significant spatial requirements, large carnivores feature prominently in many global conservation projects and policies, and the general causes of their declines are well recognised (Ripple et al. 2014, Trouwborst et al. 2017). However, this understanding has not always translated into adequate conservation action (Ripple et al. 2016). ...
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• Globally, large terrestrial carnivores (Carnivora) have suffered precipitous declines in population and range. Today, they must persist in increasingly isolated natural habitat patches within a human-dominated matrix. Effective conservation aimed at supporting carnivores in such landscapes requires species-specific understanding of habitat requirements. • We present results from a review of the published literature to assess the current state of knowledge regarding habitat preferences of the African lion Panthera leo, with the aim of identifying common drivers of habitat use across contexts. • Using the Web of Science, we identified 154 usable articles and extracted information relating to study topic, location, habitats described, land-use type, and any documented habitat preferences. • Only 31 studies documented evidence of habitat use, and collectively, they suggested that preferences for specific habitat types were varied and context-specific. The importance of prey abundance and proximity to water was highlighted in multiple studies. Anthropogenic factors interfered with expected patterns of habitat use. There was evident bias in study locations: 83% of the habitat-use studies were based in only three countries, and 70% were focussed on protected or managed areas. • Our synthesis suggests that lions demonstrate behavioural plasticity in habitat use in response to anthropogenic pressures. To understand the limits of this plasticity and to manage Africa’s changing landscapes effectively for roaming lions, future research should be focussed on analysis of habitat use outside protected areas, taking into account gradients of distance to water, prey abundance, and anthropogenic risk.
... The proliferation of border fences (a consequence of geopolitical forces such as the US-Mexico border) and contrasting protection statuses (a reflection of scalar mismatch and differing conservation priorities) exacerbates threatening processes including entanglement, encouraging harvesting and habitat conversion, and disrupting metapopulation dynamics 10,12,13 . Despite this, policy development and management actions almost always remain a matter of national sovereignty, whether crossboundary protective measures for species are warranted or not 14,15 . ...
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Rapid biodiversity loss has prompted global action to prevent further declines, yet coordinated conservation action among nations remains elusive. As a result, species with ranges that span international borders—which include 53.8% of terrestrial birds, mammals and amphibians—are in increasing peril through uncoordinated management and artificial barriers to human movement, such as border fences. Transboundary conservation initiatives represent a unique opportunity to better protect species through coordinated management across national borders. Using metrics of governance, collaboration and human pressure, we provide an index of transboundary conservation feasibility to assess global opportunities and challenges for different nations. While the transboundary conservation potential of securing multinational threatened species varied substantially, there are distinct opportunities in South-East Asia, Northern Europe, North America and South America. But to successfully avert the loss of transboundary species, the global community must be prepared to invest in some regions facing greater implementation challenges, including the nations of Central Africa, where efforts may necessitate establishing rapid conservation interventions postconflict that align with local socio-cultural opportunities and constraints. Sanctioned and coordinated approaches towards managing transboundary species are now essential to prevent further declines of many endangered species, and global policy efforts must do more to produce and enact legitimate mechanisms for collaborative action in conservation.
... CMS listing decisions tend to consider CITEScompatibility. This explains, for instance, why several cheetah populations were excluded from the species' CMS Appendix I listing ( Trouwborst et al., 2017). The CMS's influence on CITES decisions is less obvious. ...
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The relevance of the Convention on Migratory Species (CMS) for large carnivores is on the increase. Its appendices currently feature polar bear, Gobi bear, African wild dog, lion, leopard, snow leopard, and cheetah. This increased involvement raises various issues and debates concerning, inter alia, the value added by the CMS as compared to other treaties; the scope of the CMS in relation to its definition of "migratory species"; and the Convention's implications for the sustainable use of listed large carnivores. We present these and similar emerging questions within their broader context, provide beginnings of answers, and outline an agenda for further research. We further highlight the need for improved interpretive guidance on aspects of the Convention's legal text and its implications for sustainable use.
... Accordingly, the governments of the member countries (Parties) to the Convention play a role in delineating illegality of wildlife trade at an international level through their collective decisions. However, the effectiveness and universal legitimacy of CITES has been questioned (10,11), most notably in relation to large charismatic mammal species such as rhinos and elephants (12, 13). ...
Preprint
This is the final version of the previously uploaded draft entitled 'IWT: Patterns, Processes and Governance' (which contains errors). Please contact me if you would like access to a pre-print copy of this final version.
... Accordingly, the governments of the member countries (Parties) to the Convention play a role in delineating illegality of wildlife trade at an international level through their collective decisions. However, the effectiveness and universal legitimacy of CITES has been questioned (10,11), most notably in relation to large charismatic mammal species such as rhinos and elephants (12,13). ...
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... An RDH perspective allows us to shift focus and ask what units are large enough (i.e., are as small as necessary)-where the definition of necessity might be, analogous to RDH, the smallest area that enables critical ecological processes of that ecosystem or community to function, and yet within which multiple individuals (and indeed multiple species) can share. This concept might be de facto embodied in, for example, the Convention of Migratory Species that recognises that protecting the Arctic tern in its northern breeding grounds is not much use unless it is also protected in its Antarctic non-breeding grounds, or indeed that it has a right of passage between them (recognition directly relevant to Dynamo the lion, Trouwborst et al. 2017;Hodgetts et al. 2018). It is also similar to the conservation concept of metapopulation management pioneered for African wild dogs (Davies-Mostert et al. 2009), which extends the notion of a population that functions fully only by accommodating the blinking in and out of existence of a constellation of component parts. ...
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Humans, like other animals, have aptitudes and limitations shaped by natural selection; knowledge of these- A long with that of how mammals, including people, form societies and cooperate-can be fruitfully applied to the challenge of biodiversity conservation. Here, we outline the natural principles that might help conservationists (and indeed any other group concerned with political problems) get the best outcomes by aligning efforts with the grain of nature, rather than against it; we call this approach 'Natural Governance'. We illustrate the value of this perspective with reference to two aspects of conservation strategy-rethinking the spatial scales at which it is effective to plan conservation and energising human society to care enough to enable it. A Natural Governance approach stands on three critical pillars: (1) ecology (the dynamic balance between organisms and their environment), (2) cooperation (adaptations to act collectively even where this incurs short-term costs to self-interest), and (3) cultural systems (how these adaptations are manifested and vary across societies). It is commonly assumed that human society stems purely from political or socially-constructed influences. The biological perspective does not say that these influences are unimportant, but it reveals viewing humanity out of the evolutionary context to be a cripplingly narrow picture.
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The African lion is in decline across its range, and consumptive utilisation and trade of their body parts and skins has been postulated as a cause for concern. We undertook a pan-African questionnaire and literature survey to document informed opinion and evidence for the occurrence of domestic and international trade and consumption in African lion body parts across current and former range states. Sixty-five people from 18 countries participated in the online questionnaire survey (run from July 2014 to May 2015), with information provided for 28 countries (including 20 out of 24 countries believed to have extant populations). Respondents were experts within their professional spheres, and 77% had ≥6 years relevant experience within lion conservation or allied wildlife matters. Their opinions revealed wide sub-regional differences in consumptive use, drivers of trade, and access to lions that impact wild lion populations in different ways. Traditional medicine practices (African and Asian) were perceived to be the main uses to which lion body parts and bones are put domestically and traded internationally, and there is reason for concern about persistent imports from former lion range states (mainly in West Africa) for parts for this purpose. The domestic, rather than international, trade in lion body parts was perceived to be a bigger threat to wild lion populations. Parts such as skin, claws, teeth and bones are thought to be in most demand across the continent. The impact of international trade on wild populations was acknowledged to be largely unknown, but occasionally was judged to be ‘high’, and therefore vigilance is needed to monitor emerging detrimental impacts. Seventeen countries were nominated as priorities for immediate monitoring, including: South Africa, Tanzania, Zimbabwe, Mozambique, Zambia, Botswana, Kenya, Nigeria, and Cameroon. Reasons for their selection include: prevalence of trophy hunting, ‘hot spots’ for poaching, active domestic trade in lion body parts, trade in curios for the tourist market, and histories of legal-illegal wildlife trade. This survey, and increased incident reports since mid-2015 of lion poisoning and poaching in Mozambique, Zimbabwe and South Africa, and sporadic poaching events in Uganda and Tanzania, are signalling an escalating trend in the trade of lion products that is an increasing threat to some national populations. The evidence is sufficient to make more detailed investigation of this trade a conservation priority.
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The African lion is the only big cat listed on CITES Appendix II, and the only one for which international commercial trade is legal under CITES. The trade in lion body parts, and especially the contentious trade in bones from South Africa to Asia, has raised concerns spanning continents and cultures. Debates were amplified at the 2016 CITES Conference of the Parties (CoP17) when a proposal to up-list lions to Appendix I was not supported and a compromise to keep them on Appendix II, with a bone trade quota for South Africa, was reached instead. CoP17 underscored a need for further information on the lion bone trade and the consequences for lions across the continent. Legal international trade in bones to Asia, allegedly to supply the substitute ‘tiger bone’ market, began in South Africa in February 2008 when the first CITES permits were issued. It was initially unclear the degree to which bones were sourced from captive-origin lions, and whether trade was a threat to wild lion populations. Our original assessment of the legal CITES-permitted lion bone trade from South Africa to East-Southeast Asia was for the period 2008–2011 (published 2015). In this paper, we consolidate new information that has become available for 2012–2016, including CITES reports from other African countries, and data on actual exports for three years to 2016 supplied by a freight forwarding company. Thus, we update the figures on the legal trade in lion bones from Africa to East-Southeast Asia in the period 2008–2016. We also contextualise the basis for global concerns by reviewing the history of the trade and its relation to tigers, poaching and wildlife trafficking. CITES permits issued to export bones escalated from ±314y⁻¹ skeletons from 2008–2011, to ±1312y⁻¹ skeletons from 2013–2015. South Africa was the only legal exporter of bones to Asia until 2013 when Namibia issued permits to export skeletons to Vietnam. While CITES permits to export ±5363 skeletons from Africa to Asia from 2008–2015 were issued (99.1% from South Africa; 0.7% from Namibia) (51% for Laos), actual exports were less than stated on the permits. However, information on actual exports from 2014–2016 indicated that >3400 skeletons were exported in that period. In total, >6000 skeletons weighing no less than 70 tonnes have been shipped to East-Southeast Asia since 2008. Since few wild lions are hunted and poached within South African protected areas, skeletons for the legal trade appear to be derived from captive bred lions. However, confirmation of a 116kg shipment from Uganda to Laos, and reports of lion poaching in neighbouring countries, indicate that urgent proactive monitoring and evaluation of the legal and illegal trade is necessary in African lion range states where vulnerable wild lion populations are likely to be adversely affected.
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Norway's evolving national policy regarding the conservation and management of gray wolves (Canis lupus) is hotly debated. Some of the debate concerns the compatibility of this policy with the Council of Europe's 1979 Bern Convention on the Conservation of European Wildlife and Natural Habitats, to which Norway is a contracting party. This article aims to provide some further clarification in this regard, by addressing certain key dimensions of Norway's obligations under the Convention concerning wolf conservation. Specifically, it addresses the minimum population level prescribed in Article 2 of the Convention; the perspective of the transboundary wolf population shared by Norway and Sweden; and the role of the Bern Convention's Standing Committee and Secretariat. It does so by viewing relevant Convention provisions in light of public international law's basic rules of treaty interpretation, and applying those provisions to Norway's wolf policy. This analysis strongly suggests that certain basic tenets of Norway's past and current wolf policy are at odds with the country's obligations under the Bern Convention. In particular, the current wolf population target of four to six reproductions per year - a target which functions as a minimum and a maximum - appears to be well below the level required by Article 2 of the Convention.
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Many conservation professionals are familiar with the Convention on International Trade in Endangered Species (CITES), the Convention on Migratory Species (CMS), the Convention on Biological Diversity (CBD), the Ramsar Convention, and the World Heritage Convention. Regional instruments, such as those focusing on Africa, Antarctica, or Europe, are also conspicuous features of the conservation arena. Other international wildlife agreements focus on particular species, such as polar bears or albatrosses, or particular transboundary protected areas, such as the huge Kavango-Zambezi Transfrontier Conservation Area (see table 1). These agreements are collectively known as international wildlife law (Bowman et al. 2010). The binding agreements themselves are typically accompanied and informed by an evolving set of nonbinding instruments, such as Conference of the Parties (COP) decisions and action plans.
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The role of international legal instruments in biodiversity conservation has until now received relatively little attention in the conservation literature. Nevertheless, with their long-term, legally binding commitments on a transboundary scale, such instruments can be important, sometimes indispensable, implements in the conservation toolbox. After providing an overview of relevant instruments, we explore why international wildlife law matters, and what can and cannot be expected of it. We identify, on the one hand, the many different ways in which international legal instruments can deliver, and have delivered, conservation outcomes. On the other hand, we discuss the inherent limitations of international law, and provide illustrations of past achievements as well as failures. We conclude that it is worthwhile to continue to invest in making the most of international wildlife law for conservation by following an informed, selective approach. To that end, we issue a call for increased cooperation between international wildlife lawyers and other conservation professionals. There is much to be gained, partly by enhancing the legal framework itself, but especially by seizing the many opportunities for advancing the effective application of the law as it stands.
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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.
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The mounting threats posed to the global environment by harmful human activities cannot be averted without effective legislation controlling those activities. However, the environmental laws designed for this purpose are themselves under global attack. Because it is binding and enforceable, legislation is a unique and essential instrument in the overall effort to keep humanity’s impacts on the planet from transgressing critical thresholds. For instance, biodiversity laws do so by designating and protecting natural areas and controlling the exploitation of wildlife populations. Yet, due to short-term economic and other interests, such laws face constant pressures aimed at weakening their regulating impact on human activities. This new study reveals and illustrates the staggering number and diversity of tactics used to weaken biodiversity legislation across the globe. This ‘taxonomy of tactics’ encompasses dozens of categories, ranging from the creative re-definition of terms to the ‘fast-tracking’ of environmentally harmful projects, and from limiting concerned citizens’ access to court, to the silent or even express refusal of appointed authorities to enforce biodiversity laws. Whereas the predicament of the planet’s wild fauna and flora would have been even worse without the legal protection they have received so far, the onslaught against biodiversity laws has prevented these from fully performing their assigned function. The global acceleration of wildlife population declines bears witness to this. To stem the tide, strategic approaches are needed to anticipate and counter attacks on biodiversity legislation; to make the most of existing laws, including through litigation if need be; and to develop new or improved laws where necessary.
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This important and timely book provides a rigorous overview of the defining issues presently facing conservation at international level. The author provides detailed coverage of topics ranging from the classification of species right through to access and benefit sharing, drawing on his personal experience at intergovernmental level. Each question is examined through the prism of dozens of treaties and hundreds of decisions and resolutions of the key multilateral regimes, and the law in each area is supplemented by the necessary considerations of science, politics and philosophy - providing much-needed context for the reader.
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Defaunation, the emptying of ecosystems of fauna, has been highlighted as a likely threat to the conservation of carnivores, but the magnitude of this threat has yet to be quantified. We quantify the potential threat defaunation presents to wild felids. Global. For the 32 wild felids that feed primarily on mammals, we used 5,330 prey records from 237 published sources to compile a new diet dataset, FelidDIET. This dataset was used to determine the relative importance of mammalian species as prey for each felid. These data were used to quantify the relationship between felid and prey species-richness, and to estimate the potential threat to wild felids from the loss of their prey. Our analyses reveal that models that include adjusted prey species-richness as a predictor of felid-richness outperform those with less precise measures of prey-richness (potential prey-richness and total mammal-richness). This is true both when examined collectively and when split into those felids that prey upon large-bodied prey and those that prey upon small-bodied prey. For seven felid species, including six large felids (over 15 kg), 33% or more of their primary prey-species are threatened. Of most concern is the Sunda clouded leopard Neofelis diardi, for which 66.0% of its primary prey-species are threatened. In total, 57.6% of large felids’ primary prey-species are threatened or declining, compared with 26.5% for small felids. Large felids are particularly vulnerable to primary prey decline in Indo-Malaya and East and Central Africa. Our findings indicate that imminent prey loss is likely to have substantial negative effects on large felids, many of which are already highly threatened. Considering the trophic cascades associated with large predators, the threat to large felids through the loss of prey diversity presents an ecosystem-scale threat.
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Outbreeding ideas for conservation success - Volume 51 Issue 2 - David W. Macdonald, Guillaume Chapron