Leopard 2019 Leopard Trophy Hunting Quota -Wildlife Animal Protection Forum Submission
18th June 2019




18 JUNE 2019

WAPFSA Leopard Trophy Hunting Quota Submission, 2019



WAPFSA Leopard Trophy Hunting Quota Submission, 2019

WAPFSA Recommendation

The Wild Animal Protection Forum of South Africa (WAPFSA) recommends a zero quota and a cessation of the issuance of leopard trophy permits based on the available science and deficiencies of the regulatory environmental and consultative processes that are in place in the determination of population viability and status of the species.

In response to the Honourable Minister’s ‘request for information to be considered for the determination of the 2019 leopard trophy hunting quota (closing date 18 June 2019) and invitation to attend stakeholder meeting on 4 June 2019’, this submission coheres with sections 61 and 62 of the National Environmental Management: Biodiversity Act (Act No. 10 of 2004) NEMBA, in which the Scientific Authority is required to advise the Minister of the Department of Environmental Affairs on the 2019 export quota for trade in the threatened or protected species inclusive of leopards, including trophies, bones, bone pieces, bone produces, claws, skeletons, skulls and teeth of leopards for commercial purposes, which have been derived from trophy hunting in South Africa.

We, the undersigned organizations, submit that, in light of the current lack of scientific, peer- reviewed, methodologically rigorous analyses and inclusive consultative process to demonstrate beyond reasonable doubt that a non-zero export quota in trophy hunting of leopards will not imperil wild leopard survival the quota should be set to zero.

The Wild Animal Protection Forum of South Africa (WAPFSA), is an alliance of 25 diverse non- profit organizations all concerned with the welfare and protection of wild animals. WAPFSA comprises a body of considerable expertise from scientific, conservation, legal, welfare, rights, social justice, faith and public advocacy sectors.

The members of WAPFSA are:

Animal Law Reform South Africa Baboon Matters
Ban Animal Trading
Beauty Without Cruelty (South Africa) Captured in Africa Foundation
Centre for Animal Rehabilitation and Education Elephant Reintegration Trust
EMS Foundation
Elephant Specialist Advisory Group


WAPFSA Leopard Trophy Hunting Quota Submission, 2019

Four Paws (SA)
Future 4 Wildlife
Global March for Elephants and Rhinos
Global White Lion Trust
Green Girls in Africa
Humane Society International (Africa)
Institute for Critical Animal Studies (Africa)
Landmark Foundation
National Council of SPCAs
Outraged SA Citizens against Rhino Poaching (OSCAP) Southern African Faith Communities Environment Institute (SAFCEI Southern African Fight for Rhinos
Vervet Monkey Foundation
WildAid Southern Africa
Wild Law Institute


WAPFSA Leopard Trophy Hunting Quota Submission, 2019

Basis for the Zero Quota Recommendation

The leopard trophy hunting quota should be set at zero.

Globally leopard range has diminished to 25-37% of its former range and this contraction is far greater than previously known (Jacobsen et al., 2016).

In South Africa, leopard populations are currently contracting at 11% per annum (Mann et al., 2018) and a further 8% decline assessed in 2017. This indicates a persistent decline (DEA CITES Submission, 2018).

  1. A declining population of leopard is suggested by SANBI: “Results from our latest year of camera trapping paint a bleak picture of leopard population status in South Africa. Most monitored populations are declining … and the few that are stable appear to be well below their potential ecological capacity… Indeed, leopards appear to have been extirpated – at least functionally – from several sites across the country… Averaged weighted annual population growth (lambda) across all sites in South Africa was 0.89, suggesting an average population decline of 11% per year. (An 8 % decline was reported in the preceding year, 2017.) It is also important to note that we purposefully selected sites which comprise suitable leopard habitat; the situation in more marginal areas, which constitute a large part of leopard range in South Africa is likely even more dire. After more than two years without leopard trophy hunting in South Africa, there is little evidence to show that this alone is sufficient to stimulate a broad scale improvement in leopard population status.” Based on this alone any contemplation of reinstating leopard trophy hunting should be rejected. There is also evidence of preceding a persistent decline in Northern Natal (Hluhluwe-iMfolozi and St Lucia Eastern Shores) in 2015 and 2016, with the former showing a decline of 70% in five years. Further declines are reported in Limpopo with areas indicating a decline of as much as 44% (DEA, CITES Submission, 2018). Despite this, the DEA approved trophy hunting permits in Limpopo and Kwazulu Natal in 2018. By their own admission, DEA has no idea on leopard population dynamics in other regions in South Africa. Based on the trends it must be accepted that population numbers continue to be in declining across the country. The DEA request to continue the CITES allocation of 150 trophies per annum, to allow them the flexibility to issue permits up to this number, flies in the face of the data before them on population trends.
  2. Unknown numbers of leopard mortalities from human-wildlife conflict and “cultural” artefact harvesting compounds this situation and should advocate against the reinstitution of leopard sports hunting. Government has no control over the situation and should not be compounding the peril for the species by contributing to its mortalities in appeasement of the hunting industry pressures.
  3. Regional genetic and habitat fragmentation has only been demonstrated in the Eastern and Western Cape. It may be evident in other areas of population fragmentation and restricted connectivity. Until this is investigated further, removal of individual leopards may in fact contribute significantly to the demise of the species


WAPFSA Leopard Trophy Hunting Quota Submission, 2019

through ignorant actions. This should be researched thoroughly prior to any contemplation of reinstitution of leopard hunting.

  1. Flawed monitoring by SANBI, which they claim themselves rather robustly is the “gold standard”, is rudimentary and flawed. It is based on a method that cannot remotely establish asymptote (within 30 – 60 days of camera trapping) of population estimates (of which a closed population requires 90 days (Kamaranth. 2009)). This fails to consider social dynamics and qualitative analysis of population structures and dynamics and pays no regard to the genetic structure of populations. Trophy hunting can exacerbate population declines where other human-caused mortality is severe, ongoing, not fully recorded and uncontrolled.
  2. Improper consultative process has a long and ingrained culture in terms of leopard conservation, threatened and protected species legislation and damage causing animal legislative issues with Department of Environmental Affairs (DEA) and the South African National Biodiversity Institute (SANBI). It dates to the Leopard PHVA in 2005 when stakeholders were excluded from the process, which had almost no detailed data, on which it was based and the subsequent CITES permit allocation number of 150 trophies per annum. The latter has never been rationally reviewed, and by DEA’s own version was retained for no other reason than the onerous process of changing trophy allocation limits with CITES. Even if the science dictates that current allocations are unsustainable, this improper consultation continued as detailed below with a recent trend where a select few individuals who play “prosecutor, judge and jury” on a consultative process and who are appointed without due government processes with nepotistic relations and biased views supporting continued trophy hunting. This grouping lead by SANBI personnel have drafted the Threatened and Protected Species regulations, the non-detriment findings report for CITES (NDF), the Norms and Standards for hunting leopards, recommendations for instituting leopard hunting in 2018, the misrepresented CITES submission of 2018, and now control advisory process to this attempt to recommence the hunting of leopards. DEA has failed court processes due to this manner of consultative processes in the lion hunting and rhino horn trade cases and it seems that nothing has been learnt from these flawed consultative processes. DEA has participated in misrepresentation of information to CITES in 2018 in its request to retain the 150 trophy permits in 2018 ahead of the CITES meeting this year. It has failed to cite relevant, current published research while biasedly presenting conjecture that supports the continued hunting of leopards in the face of data that points to continued decimation of the leopard regional populations. The continued motivation of trophy hunting flies in the face of the available science, and the very principles that the Department claims to be adhering to, namely:a. Sustainable use paradigm – even if this was adhered to the decreasing population, other human caused mortalities and continued habitat fragmentation, trends points to current unsustainable management;

b. Adaptive management is not adhered to in the face of SANBI’s own data, showing an 11% population shrinking in 2018, based on severely depleted and fragmented leopard populations


WAPFSA Leopard Trophy Hunting Quota Submission, 2019

c. “Involvement of stakeholders” belies SANBI and DEA practices by excluding stakeholders, and further concealing data which is evident in the declining leopard population figures from 2017 and 2018;

  1. Robust science is being ignored in reports and not adhered to in monitoring;
  2. The impact of damage causing animal deaths and illegal killing is unverified, not accurately recorded and seemingly ignored in quota application andissuances;
  3. There is no basis in science that support the use of sports hunting in themanagement of sustainable leopard populations;
  4. There is no scientific basis whereon sports hunting of leopards has provenbeneficial for local livelihoods.
  1. The current farcical approach of approving permits on a baseline that is set arbitrarily on the commencement of the current SANBI initial monitoring results of 2016, has no basis on historical carrying capacity or appropriate natural distribution and population densities.
  2. NDF emerged with a negative finding for leopards and as such should dictate zero quota allocation in terms of CITES until a positive NDF replaces it. Despite this, the DEA and SANBI, asserted that permits should be issued at the allocated 150 per annum rate in the NDF report and their current submission before CITES as submitted in 2018. After a public outcry following the NDF report, the DEA, correctly, set the quotas at zero for two years (2016 and 2017) and yet in 2018, despite the standing negative finding on the NDF reinstituted leopard hunting quotas without due consultation and scientific justification and in spite of population declines evident from their “monitoring” project results. All attempts at transparency were resisted and when the SANBI report of 2018 detailed the contraction of 11% in the population became apparent through the leaking of the report by some parties, the reason for the non- accountability became obvious, as there appeared to be no factual basis for the recommencement of trophy hunting in the face of the continued population declines.

Therefore, based on 1) the declining population, 2) the escalation and ongoing threats to the population, and 3) the poor data quality and 4) undue process in consultation, it is incumbent on the DEA to decline any current permits for trophy hunting or harvesting of the species. We recommend that a zero quota be adopted and that this should stay in place until appropriate, accountable and transparent governance processes are employed, a NDF be confirmed and that decisions are based on respected scientific methods, and not the current sham processes.

No leopard trophy hunting quotas should be allocated, until a positive NDF is adopted and therefore should be in accordance with CITES approvals of a zero quota and not as present attempts to wrongfully keep this allocation in the NDF and in the 2018 proposal to CITES at 150.


WAPFSA Leopard Trophy Hunting Quota Submission, 2019

Original Quota of 150

The original quota of 150 CITES permits stems from the PHVA of 2005 and has never been reviewed. The 2005 process was notable by the absence of any credible data on which to base the quota amount and the exclusion of key stakeholders from the process. By its own admission, DEA in their submission to CITES in mid-2018, retained the quota application at 150 for no rational scientific-based reasoning, and by its own admission retained it at 150 to avoid applying for changed quota quantum as it would be administratively too onerous to do such an application, even if the science dictated a reduced quota was warranted.

This together with the assertion that it should retain the 150 quota in the face of a negative NDF points to irrational and nefarious agendas to continue trophy hunting of leopards regardless of the objective facts before them.

Flawed Monitoring Processes

Focus on quantitative assessment of leopard populations, almost exclusively in protected areas, fails to assess leopards in production landscapes where the conflict zones exist. Any results obtained from protected areas may in fact be an over-estimation of leopard populations when extrapolated beyond these protected areas. The method fails to follow recommended scientific methods of establishing asymptote measures to obtain a population estimate using camera traps by only placing these monitoring cameras at sites for 30 – 60 days instead of the recommended 3 months (Ref. Karanth, 2009).

The monitoring process fails to do any meaningful population dynamic analysis and assessments of social structures of leopards and the impacts mortalities have on these. There is also no investigation on population connectivity in transformed landscapes. No investigation of population genetic structure is done and the impact this has on genetic vitality of the population in areas of human-caused mortality, and thus on population persistence. Furthermore the SANBI team self-grandiosity by claiming their methods are the “gold standard” fails to accept the fatal flaw that their measure of the starting assessment of 2016 is the basis from which recommendations were made whether populations were increasing or decreasing, when already this starting point was substantially reduced from natural carrying capacities. The basing of permit allocations on this starting point was based on a flawed assumption. These flaws in monitoring makes a mockery of the claimed adaptive management underpinning permit recommendations and the science that ought to underpin such methods.


WAPFSA Leopard Trophy Hunting Quota Submission, 2019

Government Legal Obligations and Consultative Process Failures

Much frustration has been experiences as stakeholders over almost 15 years of attempts to get accountable governance in government processes affecting leopard conservation involving the following:

1. Leopard PHVA assessment of 2005
2. Norms and standards for the management of Damage Causing Animals, 2010
3. Threatened or Protected Species Regulations, 2013
4. Non-Detriment Findings on leopards, 2015
5. Protesting of the appointment of Panthera as specialist consultants to SANBI without

due process
6. Norms and standards for the management of Damage Causing Animals, 2016
7. Draft Norms and Standards for the Management and Monitoring of the Hunting of

Leopard in South Africa for Trophy Hunting Purposes, 2017
8. Reinstating of leopard trophy hunting permits in the presence of a negative NDF in

2018 and refusal to provide the scientific justification for such.
9. The application to CITES in 2018 to retain the allocation of 150 CITES permits in the

presence of a negative NDF and claiming their application is based on robust science and accountable governance when this was untrue.

This has been mostly centered around a select unaccountable group of state officials and a few consultants appointed without due process affecting leopard management decisions, policy and legislation development and subsequent permit issuing that have been unaccountable, based on biased and jury-selected facts and ignoring the science that dictates the need to set the hunting quota at zero.

It is recommended that accountable governance structures be developed to develop leopard conservation policies, legislation and administrative processes that are defendable scientifically and that individuals that have developed a cabal around leopard conservation in the past be excluded from such processes.

Approach to be adopted in relation to determination of detriment

The Convention on International Trade in Endangered Species (“CITES”) provides that an export permit for a specimen of a species included in Appendix II cannot be granted unless the Scientific Authority of the State of export has advised that such export will not be detrimental to the survival of that species. Species are included in the Appendices to CITES if they are, or may be, detrimentally affected by international trade. Consequently, it must be assumed that international trade has a detrimental effect on any listed species unless the Scientific Authority determine that it will not, by making a so-called “non-detrimental finding ” (“NDF”).

In other words, an NDF is an exception to the general rule that prohibits trade in CITES Appendix II-listed species and the Scientific Authority may only make a NDF if there is clear scientific evidence that trade will not have any detrimental effect. If the available evidence is not conclusive in this regard, the Scientific Authority cannot, and must not, make an NDF. This is consistent with the principle in the National Environmental Management Act (“NEMA”)


WAPFSA Leopard Trophy Hunting Quota Submission, 2019

which requires that the decision maker must apply “a risk averse and cautious approach … which takes into account the limits of current knowledge about the consequences of decisions and actions” (NEMA, section 2 (4)(a)(vii)). Put differently, in the absence of sound scientific justification for a specific quota, the Scientific Authority must not set a quota.

Proposals to establish or amend an NDF must evidence-based and include details of the scientific basis for any quota that may be proposed. CITES does not specify precisely what scientific evidence is required but the CITES conference of the parties has adopted guidelines in relation to species such as sharks. In 2002 the IUCN published a checklist to assist scientific authorities in making non-detriment findings for Appendix II exports. The checklist helps Scientific Authorities to identify the factors that need to be taken into account when making an NDF and the strengths and weaknesses of the available information. (At a minimum the Scientific Authority should consider information such as: species distribution, population status, population trends, threats, utilization and trade, actual or potential trade impacts, population monitoring, management and control measures.)

South African Scientific Authority must take account of all relevant information

Despite the existence of these (non-binding) guidelines, when deciding whether or not to make a NDF the Scientific Authority must act both in a manner that is not contrary to CITES and in accordance with the law that governs it (in this case South African law). As explained below, South African law requires the Scientific Authority to take account of all relevant information, not just information from peer-reviewed scientific studies or from scientists.

The decision by the South African Scientific Authority about whether or not international trade may be detrimental to leopards, is a decision that may significantly affect the environment. Consequently in making that decision the Scientific Authority must be guided by the environmental right in section 24 of the Constitution and by the national environmental management principles in section 2 of the National Environmental Management Act (“NEMA”).

The following principles are particularly relevant to this decision:

2(4)(a)(vii) that a risk averse and cautious approach is applied, which takes into account thelimits of current knowledge about the consequences of decisions and actions; and

(b) Environmental management must be integrated, acknowledging that all elements of the environment are linked and interrelated, and it must take into account the effects of decisions on all aspects of the environment and all people in the environment by pursuing the selection of the best practicable environmental option.

(g) Decisions must take into account the interests, needs and values of all interested and affected parties, and this includes recognizing all forms of knowledge, including traditional and ordinary knowledge.
(o) The environment is held in public trust for the people, the beneficial use of environmental resources must serve the public interest and the environment must be protected as the people’s common heritage.”

A decision to establish or amend a quota for the export of leopard trophy hunts would also be “administrative action” for the purposes of the Promotion of Administrative Justice Act (“PAJA”). Consequently, the decision-making process must be procedurally fair, and the


WAPFSA Leopard Trophy Hunting Quota Submission, 2019

decision-maker must, among other matters, take all relevant considerations into account, and disregard irrelevant considerations.

Not only is a negative NDF in place for leopard trophy hunting and exports, but there are substantive problems with the governance around decision making related to leopard conservation, yet the DEA have continued and restarted trophy hunting and continued to request a maintenance of 150 CITES permits for leopards when the data and science does not support such.

Consideration of the effect of a leopard trophy hunting and export quota on the survival of all wild leopards
It is important to note that the Scientific Authority must consider the impact of a leopard trophy hunting and export quota on leopards both within South Africa and in other range states. Although CITES defines “species” as “any species, subspecies, or geographically separate population thereof”, the 10th meeting of the Conference of the Parties to CITES adopted a resolution on the Designation and role of the Scientific Authorities (resolution Conf. 10.3) which recommends that:

“the appropriate Scientific Authority monitor the status of native Appendix-II species and export data, and recommend, if necessary, suitable remedial measures to limit the export of specimens in order to maintain each species throughout its range at a level consistent with its role in the ecosystem and well above the level at which the species might become eligible for inclusion in Appendix I;”(2 j – emphasis added)

This means that in order to make an NDF the decision-maker must not only have adequate scientific evidence that trade in leopards is not resulting in, and will not result in, higher mortalities among wild leopards in South Africa, it must also have adequate scientific evidence to exclude the risk that permitting a legal trade in leopards will not be detrimental to the prospects of maintaining appropriate leopard populations in all range states. Establishing a leopard hunting industry is detrimental to the long-term survival of the species in the wild. The Scientific Authority cannot make a NDF in order to set a quota for the export of leopards unless it has the evidence to prove the contrary.

The 2018 quota was not established on scientific grounds

As the call for submissions rightly points out, the Scientific Authority is required to make recommendations to the Minister ‘based on a scientific and professional review of all available information.’ We submit that this was in fact not complied with when the new quota was set in 2018. Such a review would have revealed that no non-detriment finding (NDF) pertaining to the trade in leopard trophies (an Appendix II-listed species according to the Convention on International Trade in Endangered Wild Species of Fauna and Flora (CITES)) and its potential impact on wild leopards yet existed. The 2015 NDF carried out to assess whether a trade in leopard trophies was detrimental to wild leopards in South Africa came out against trophy hunting of leopards (a negative NDF). The negative NDF is still in place. Despite this negative NDF the scientific authority tried to assert that permits still be issued and only relented after public and media pressure but in 2018 applied to maintain the allocation of 150 as it was too inconvenient to have this changed with CITES and they recommenced permit issuing. Based on the negative NDF, it would therefore appear to us to have been prudent to set the quota to zero until such time as, at a minimum, a relevant NDF

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had been issued. In the interim data emerged that pointed to accelerating population declines and the Scientific Authority and small select group of consultants tried to conceal the negative population data from stakeholders while moving ahead with reinstating trophy quotas. Such concealment can only point to bad faith and nefarious agendas at appeasing pressure from the trophy hunting industry.

The precautionary principle appears to require that the burden of proof is on the proposer of an action to demonstrate that it will have no harmful or irreversible effect. The bottom line is that a scientific review would likely have concluded that enough warning signs were present to suggest that a quota should not be established until more scientific research was available.

Scientific Justification for Recommendation

Large carnivores have experienced significant population declines due to geographic range contractions, fragmentation of their habitat and human wildlife conflict events (Morrison et al., 2007), other human caused mortality and the illegal wildlife trade. All carnivore ranges have had a significant decline globally (Ripple et al. (2014). In South Africa this is supported by the absence of almost all free roaming large carnivores, except for leopard and occasionally cheetah and wild dogs. The one remaining large felid in parts of South Africa is the leopard (Panthera pardus); but the worldwide range is continuing to decline. This global range loss is more extensive than previously appreciated and the species occupies a mere 25-37% of its historical range. Globally the decline of leopard populations is illustrated (Figure 1&2) as published in Jacobsen et al. (2016). Regionally (the whole of South Africa) the leopard is listed as Vulnerable C1 (Swanepoel et al., 2016), although the southernmost population in the Cape Provinces may be substantially more threatened. Additionally, it is highlighted that leopards are undergoing increasing levels of persecution due to being hunted in retaliatory killings, for cultural regalia and from the increase in high-value game breeding farms that treat them as damage causing animals within their current distribution. This is leading to significant and continuing declines, 10% over three generations. Even internal reports from SANBI suggest alarming declines of leopard population by as much a 11% per annum in 2018 is also recorded by the Scientific Authority, and in their own words the “results from … latest year of camera trapping paint a bleak picture of leopard population status in South Africa. Most monitored populations are declining…and the few that are stable appear to be well below their potential ecological capacity … Indeed, leopards appear to have been extirpated – at least functionally – from several sites across the country…Averaged weighted annual population growth (lambda) across all sites in South Africa was 0.89, suggesting an average population decline of 11% per year (our emphasis)!

Actions of the Scientific Authority should rather be focused on that which can facilitate the conservation of the species instead on this yearly battle to limit their further sanctioned human-caused mortality through trophy hunting. This should include:

  • Connectivity of isolated populations,
  • Management of genetic bottlenecking of remaining populations,
  • Addressing human-wildlife conflict,

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• Utilizing and evaluating efficacy of local translocation as a means of dealing with conflicts, and

• Reducing human caused mortality.

Figure 1&2. Current and historical leopard distribution demonstrating the global decline Jacobsen et al. (2016).

Population Connectivity: Devens et al. (2018) estimated the population of leopards to be between 467-533 individuals in the Western Cape and between 365-430 in the Eastern Cape. However, these free roaming leopards are facing hazards as the anthropogenic landscape expands, and concurrent habitat destruction. The above estimates may be an overestimation because it relies on extrapolated habitat patches that include small (narrow and isolated) patches that probably do not support this wide-ranging species. Such a landscape can result in movement constraints and increased mortality events resulting in a declining population (Woodroffe et al., 2005). The Northern, Western & Eastern Cape have low population of leopards with which to effect conservation of the species, as part of that connectivity between isolated populations forms a key part of such conservation actions. The detailed peer-reviewed population densities in other areas of South Africa is not known and the current leopard monitoring by SANBI and Panthera fail to meet scientific muster as the period of monitoring fails to establish asymptote for population estimates. In the absence of this detailed scientific evidence of acceptable and healthy populations there is no basis to move ahead with authorizing trophy hunting to add to the already accepted evidence of increasing human caused mortality and perilous nature of leopard populations globally and regionally.

Drouilly et al. (2017) performed an analysis of carnivore scats found both in protected areas and on farmland. They found that leopards located in protected environments had no evidence for livestock predation events in their scats, and preferentially preyed on antelopes

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and bushpigs (although their study sites were not representative of leopard habitat across the range of leopards). No leopard scat was detected on any of the farmlands on their particular farming site; reinforcing the knowledge that these anthropogenic regions form considerable barriers to leopard movements. This can ultimately lead to a genetic bottleneck (Sweanor et al., 2000). A consideration of apex predators should form a crucial component of conservation strategies and should be considered on an equal footing to habitat preservation efforts.

Genetic Bottlenecking: McManus et al. (2015) studied the genetic structure of leopards in the Eastern and Western Cape. They found that three distinct population groups existed within the study area. Two of these populations in the north and south of the Western Cape were found to be acting as sink populations with a low rate of gene flow. They suggested that this population structure is a result of increased anthropogenic land use and human wildlife conflict events. McManus (PhD Thesis, 2018) has additionally shown that both male and female leopard behaviours are strongly influenced by their distance to roads and towns. They seldom use human modified landscape (ploughed fields, orchids, etc.) supporting the claims that the modified environment is causing the low gene flow and the bottlenecking of the populations of leopards in the Western Cape. Additionally, it was found that males organized themselves based on the locations of females, whilst females made few inter-patch movements when close to towns. This suggests that leopards surrounded by highly modified environments will find themselves increasingly isolated with only a few males making large dispersal movements across the anthropogenic landscape. McManus (PhD, 2018) detailed resource usage of leopards in the Eastern and Western Capes and illustrated the habitat and population fragmentation of the species in the study area. Furthermore, the projected corridors of population connectivity were identified.

Very few areas in South Africa has had populations studied to this detail which is needed prior to authorization of further leopard mortality actions, such as trophy hunting authorization.

Very little scientific work has been done to evaluate population structures of leopard across the country. In the light of range shrinkage, ongoing population and habitat fragmentation and mortalities it becomes more and more important to investigate population and genetic structure of leopard populations before authorizations for further human mortalities. McManus et al. (2015) showed that the leopards of the Eastern and Western Cape form a complex genetic structure with low rates of gene flow between three identified core subpopulations. This suggest genetic bottlenecking of isolated and fragmented leopard sub- populations. Habitat fragmentation is suggested as the primary cause for the reduction in rates of gene flow between these populations which is resulting in a significant genetic bottle- necking effect on the southern and northern populations in the Western Cape (McManus et al., 2015). Establishing connectivity between populations and reducing further human caused mortality is crucial to avoid any further genetic deterioration. An understanding of the genetic landscape will greatly help in guiding conservation orientated decisions, in particular for mortality decisions, future translocations of leopards and for the management of subpopulations and protected areas by conservation bodies, particularly supporting connectivity corridors being functional through addressing and mitigating conflict and effective gene flow and translocations in select instances. Fostering gene flow between isolated populations through territorial connectivity or active management, such as

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translocations, may be important, because deterioration of gene flow can have detrimental effects within a few generations that would result in low population survival. The absence of this detailed knowledge together with glib decision on authorizing leopard hunts may have massively detrimental effect on the genetic viability of populations in the current state of population declines and habitat fragmentation.

Addressing Human Wildlife Conflict: Leopards are difficult to study because of their elusive nature. Most studies have focused on leopards within nature reserves; areas where leopards are significantly threatened often generally overlooked (Balme et al., 2014). The current SANBI studies are flawed in that the methodology they use is unable to establish asymptote in population quantitative assessments and fails to deal with behavior studies and genetic analysis, both latter being crucial to assessing population viability. A focus on production landscapes is essential and it holds the potential to address key knowledge gaps. Current monitoring is focused on protected areas. Those leopard populations which persist in anthropogenic environments have received little scientific attention, often due to the difficultly of studying a highly elusive animal in such a landscape. As a result, conservation practices and management plans for leopards utilising these habitats are all too often based on anecdotal evidence. Balme et al. (2014) identified several threats faced by leopards in South Africa; key among these were loss and fragmentation of habitat and anthropogenic killing. It seems to be illogical to be considering trophy hunting of leopards when there is little to no veracity to any data related to mortalities from human wildlife conflict situations, and even less knowledge of cultural harvesting of leopard body parts/skins. In the face of the global decline of leopard populations, a SANBI reported annual decline of 11% in the population of leopards locally, it seems utterly dumbfounding that any consideration is given to leopard trophy hunting.

The overt threats by game farmers and hunting community that they ought to be given access to trophy hunting permits to allow them to seeing value in this species or else they may resort to “shoot, shovel and shut up” practices are nothing short of extortion. The species is a res nullius public good and permits are a state mechanism to assign such public assets to private profit and in the face of the decline in the species, this public good for their existence value outstrips the private profit of hunting outfitter or pro-hunting landowners. Published research comparing retaliatory killings with trophy hunting and its effects on leopard populations suggest that the former has greater negative impacts on leopard populations, but the study fails to consider any compensation schemes other than trophy hunting (Swanepoel et al., 2015). This is a constant failure and an indicative of a constant bias and appeasement of the trophy hunting industry even in the face of threats to the species by this bias.

Government would do well rather to consider the development of compensation schemes for human wildlife conflict situations. Wildlife compensation schemes have been found to improve tolerance of carnivores in conflict areas by raising awareness and shifting economic responsibilities from farmers to broader communities (Wagner et al., 1997). This allows animals to live longer and encourages the implementation of alternative controls rather than long-used lethal management methods. Where compensation schemes are absent, retaliatory killing is more likely (Nyhus et al., 2005). However, damage to property by wildlife will occur in perpetuity and over vast areas, thus formalizing sustainable compensation schemes has challenges. Furthermore, even individuals receiving compensation may kill

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animals illegally. It is a reality that leopards at times and opportunistically do hunt and kill livestock and indeed framed game. This has historically led to farmers taking lethal control measures to remove the problem causing animal, with these actions taking place both legally and illegally. Conservation authorities have also killed leopards that they deemed to be “problem leopards”. Such actions have not been open to review and scrutiny. However, this has the potential to create a larger problem where a territorial gap opens in the landscape causing disruption within the underlying population structure. It is believed and proposed that an effective solution is to better manage the livestock, rather than the predator, and in limited circumstances where key threatened or protected species (such as leopards) are imperiled by human wildlife conflict to provide compensation for any future livestock losses caused by leopards. It is also very important that human wildlife conflict situations NOT BE CONFLATED into trophy hunting events as all high value trophy animals will be incentivized to be labeled problem animals.

Leopard Removal and Translocations: Instead of trophy hunting we would suggest that if leopards are to be condemned to removal that translocations are rather considered as it may be a valuable conservation tool in threatened populations and species. Many leopards have been translocated or killed that have been deemed problem animals. These actions have been done without scientific analysis and review. In the scenario of low numbers, isolated sub- populations, limited connectivity, continued persecution of leopards and a population with evident reduced genetic diversity amongst extant populations, it has become important to investigate the feasibility and parameters of the success of translocations. Translocation also may be a very effective tool for rewilding efforts.

Translocation as a conservation tool has been identified by the IUCN as a critical component of conserving endangered species and in re-establishing populations in areas where species have been driven to extinction (IUCN/SSC, 2013). They recognize that reinforcement of existing populations to enhance viability, as well as reintroduction to establish new viable populations are equally vital and should be considered in tandem. Fonturbel el al. (2011) performed a meta-analysis of 50 scientific articles concerning translocations of large carnivores around the world; they found that in 64% of cases the translocated individual did not return to their home range (N=15). However, this does not equate to a 64% success rate. On the contrary 47% of the animals translocated died within 110 days, with 83% of deaths caused as a direct result of human activity, both intentional (illegal hunting) and accidental (vehicle collisions). Although many of the study species were large felids, none of the studies analysed focused on leopards. Translocation has been shown to be extremely successfully in managing the cheetah (Acinonyx jubatus) meta-population that exists on small private game reserves in South Africa (van der Merwe et al., 2016), which can be contained by fences. This network of reserves essentially forms a fragmented habit for cheetahs where no natural genetic flow takes places; without translocation each of these populations would be unviable. However, these animals enter range-restricted (cheetah proof fence) habitats allowing a high degree of success. There are moves afoot to manage both cheetah and lion meta-populations through judicious translocations.

7-year age male hunting limit: The entire hunting quota based on the purported fact that hunting male leopards over 7 years of age has minimal “impact on leopard population persistence” (Government Gazette, No 40601, 8 February 2017). Packer et al. (2009) is quoted

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as the reference for this. When the paper in question is analyses it says nothing of the sort. It further should be noted this paper uses data gleamed from CITES import data to the USA, and no empirical data and does modelling based on this. It deals with the leopard population of one protected area but fails to present the details from Phinda (also the work of Balme and Panthera). Additionally, it broadly states that other population data was used, including from a highly criticized article (Martin and Meulenaar, 1988). The study is fraught with faults and is known to be dramatically inflated and unreliable as criticized extensively by peers. Furthermore, the paper fails to detail the data and methods to justify models used to derive these conclusions, nor is there any attempt at validating the model. It must be stated that the reliance on CITES data is very suspect as there is already a “filtered” element to this data and amounts to nothing short of opinion surveys and not imperial fact. It is of grave concern that the DEA is prepared to rely on this questionable data (CITES import data), a suspect modelling exercise and opinions about the 7-year limit to base an entire national N&S of harvesting of leopards. It seems cavalier to say the least. How the conclusion can be made off this glibly used “fact” is incomprehensible? No other research has attempted to validate this claim nor confirmed it through verifiable and repeatable empirical research. The opinion of the 7-year of age threshold is presented as scientific fact of what appears to be questionable conclusions. The effect of action that could flow for this gives no regard population social structures. It cannot be accepted to be the guide for policy. We would urge DEA to review this paper critically and reconsider this erroneous assertion that male leopards can be harvested after 7 years of age without population persistence impacts. It uses unclear methods in model design and data known to be highly inflated leopard density data.

Leopards probably live to ages 10 -13 in protected environments, and when in conflict in production landscapes on average 6-8 years. It is rare that a male leopard has a stable territory before 5-6 years and thus the paternal factor of cubs. To suggest you can shoot them from this early age is sure to create havoc amongst the populations and generalize infanticide. It also gives absolutely no regard for the impact on social structures and the impact this has on species persistence. The fact remains that we know almost nothing about leopard population studies and the few detailed empirical studies that have been done are so regionally specific that making sweeping statements based of “models” and extrapolation of CITES data and a tiny regional based inquiry is irresponsible to the extreme and will not provide insight to a broader areas. We would suggest that the 7-year threshold to kill males is opinion masquerading as fact and should be rejected.

The second aspect that is fatally flawed is that leopards can’t be aged and sexed accurately to abide by the first already flawed assumption. We believe this to be a fabrication in support of the flawed assertion above. Panthera’s own research bears this out and demonstrates the poor accuracy of sex and age predictions even amongst photographic guides and researchers, and much worse amongst hunters which apparently had the least accurate assessments (Balme et al. 2012). More concerning is that this study excludes the human factor in leopard identification, as it uses still, and clear images and suggest fleeting and field sightings that can’t be used as an identification. It goes on to disprove that hunters can be trained to improve their accuracy. SANBI and DEA have accepted this without question in support of authorizing leopard trophy hunting. There is no evidence for this. The ability to accurately sex and age leopards in real life in the field is not possible, not implementable or police-able and

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thus a fanciful construct to enable hunting under false pretense of management. This is even more improbable if not impossible in the heat of a trophy hunt scenario.

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Balme, G.A., Lindsey, P.A., Swanepoel, L.H. & Hunter, L.T.B. 2014. Failure of research to address the rangewide conservation needs of large carnivores: Leopards in South Africa as a case study. Conservation Letters 7: 3–11. doi:10.1111/conl.12028

Brashares, J.S., Prugh, P.R., Stoner, C.J. & Epps, C.W. 2010. Ecological and Conservation Implications of Mesopredator Release In: Terborgh, J, Estes, J.A. (eds) Trophic Cascades: Predators, Prey, and the Changing Dynamics of Nature. Island Press, Washington, DC, pp. 221–240.

Department of Environmental Affairs. 2018. Leopard Quota Review (for CITES) AC30 Doc 15, Annex 3

Devens, C, Tshabalala, T, McManus, J. & Smuts, B. 2018. Counting the spots: The use of a spatially explicit capture recapture technique and GPS data to estimate leopard (Panthera pardus) density in the Eastern and Western Cape, South Africa. African Journal of Ecology.00:1–10. https://doi.org/10.1111/aje.12512

Drouilly, M., Nattrass, N. & O’Riain, M.J. 2017. Dietary niche relationships among predators on farmland and a protected area. Journal of Wildlife Management. doi:10.1002/jwmg.21407

Fonturbel, F.E. & Simonetti, J.A. 2011. Translocations and human-carnivore conflicts: problem solving or problem creating? Wildlife Biology 17: 217–224.

Getz, W.M., Wilmers, C.C. 2004. A local nearest-neighbour convex-hull construction of home range utilization distributions. Ecography, 27: 489-505.

Getz, W.M., Fortmann-Roe, S., Cross, P.C., Lyons, A.J., Ryan, S.J., Wilmers, C.C. 2007. LoCoH: non- parameteric kernel methods for constructing homes ranges and utilization distributions. PLoS One, 2(2): e207.

Grimbeek, A.M. 1992. The ecology of the leopard (Panthera pardus) in the Waterberg. M.Sc. thesis, University of Pretoria, Pretoria.

Gopalaswamy, A. M., Royle, J. A., Hines, E. J., Meredith, M. E., & Singh, P. (2012). Program SPACECAP: software for estimating animal density using spatially explicit capture-recapture models. Methods in Ecology and Evolution, 3, 1067–1072.

IUCN/SSC, 2013. Guidelines for Reintroductions and Other Conservation Translocations. Version 1.0. Gland, Switzerland: IUCN Species Survival Commission, viiii + 57 pp.

Jacobson, A.P., Gerngross, P., Lemeris, Jr. J.R., Schoonover, R.F., Anco, C., Breitenmoser-Würsten, C., Durant, S.M., Farhadinia, M.S., Henschel, P., Kamler, J.F., Laguardia, A., Rostro-García, S., Stein, A.B. & Dollar, L. (2016). Leopard (Panthera pardus) status, distribution, and the research efforts across its range. PeerJ 4:e1974 https://doi.org/10.7717/peerj.1974

Kie, J. G., et al. 2010. The home-range concept: are traditional estimators still relevant with modern telemetry technology? Phil. Trans. R. Soc. B, 365: 2221–2231

Labuschagne, C., Dalton, D. L., Grobler, J. P., & Kotzé, A. (2017). SNP discovery and characterisation in White Rhino (Ceratotherium simum) with application to parentage assignment. Genetics and Molecular Biology, 40(1), 84–92. http://doi.org/10.1590/1678-4685-GMB-2016-0058

Lichti, N. I., and Swthart, R. K. 2011. Estimating Utilization Distributions with Kernel Versus Local Convex Hull Methods. Journal of Wildlife Management, 75(2): 413-422.

18 | P a g e

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Lyons, A.J., Wendy, C.T., Getz, W.M. 2013. Home Range Plus: A space-time characterization of movement over real landscapes. BMC Movement Ecology.

Lyons, A.J. 2012. Visualizing Wildlife Conservation and Development in Southern Africa: A Multi- Optic Approach. Ph.D. University of California, Berkeley.

Mann, G., Pitman, R., Broadfield, J., Taylor, J., Whittington-Jones, G., Rogan, M., Dubay, S., Balme, G. 2018. South African Leopard Monitoring Project: Annual report for the South African National Biodiversity Institute (SANBI/Panthera Report)

McManus, J.S., Dalton, D.L., Kotzé, A., Smuts, B., Dickman, A., Marshal, J.P. & Keith, M. 2015. Gene flow and population structure of a solitary top carnivore in a human-dominated landscape. Ecology and Evolution 5: 335–344. doi:10.1002/ece3.1322

McManus, J., Dickman, A., Gaynor, D., Smuts, B., & Macdonald, D. (2015). Dead or alive? Comparing costs and benefits of lethal and non-lethal human–wildlife conflict mitigation on livestock farms. Oryx, 49(4), 687-695. doi:10.1017/S0030605313001610

McManus, J.S. 2018. Investigating the landscape connectivity and management of leopard (Panthera pardus) in the Eastern and Western Cape, South Africa. Ph.D. University of Witwatersrand, Submitted

Mills, M.G.L. 1991. Conservation management of large carnivores in Africa. Koedoe 34: 81–90.

Morrison, J.C., Sechrest, W., Dinerstein, E., Wilcove, D.S. & Lamoreux, J.F. 2007. Persistence of large mammal faunas as indicators of global human impacts. Journal of Mammalogy 88: 1363–1380. doi:10.1644/06-MAMM-A-124R2.1

Nyhus, P.J., Osofsky, S.A., Ferraro, P., Madden, F. & Fischer, H. 2005. Bearing the costs of human- wildlife conflict: the challenges of compensation schemes. In R. Woodroffe, S. Thirgood, & A. Rabinowitz (Eds.), People and Wildlife, Conflict or Co-existence? (Conservation Biology, pp. 107-121). Cambridge: Cambridge University Press. doi:10.1017/CBO9780511614774.002

Pool-Stanvliet, R., Duffell-Canham, A., Pence, G. & Smart, R. 2017. The Western Cape Biodiversity Spatial Plan Handbook. Stellenbosch: CapeNature.

Ripple, W.J., Estes, J.A., Beschta, R.L., Wilmers, C.C. & Ritchie, E.G., Hebblewhite, M., Berger, J., Elmhagen, B., Letnic, M., Nelson, M.P., Schmitz, O.J., W. Smith, D.W., WallacH, A.D. & Wirsing, A.J. 2014. Status and ecological effects of the world’s largest carnivores. Science 343: 6167. doi: 10.1126/science.1241484

Royle, J. A., Karanth, K. U., Gopalaswamy, A. M., & Kumar, N. S. (2009).

Bayesian inference in camera trapping studies for a class of spatial capture-recapture models. Ecology, 90, 3233–3244. https://doi.org/10.1890/08-1481.1

Sunquist, M.E. & Sunquist, F.C. 2001. Changing landscapes: consequences for carnivores In: Gittleman, J.L., Funk, S.M., Macdonald D.W. & Wayne, R.K. (eds) Carnivore Conservation. Cambridge University Press, Cambridge, UK., pp. 399–418.

Swanepoel, L.H., Balme, G., Williams, S., Power, R.J., Snyman, A., Gaigher, I., Senekal, C., Martins, Q., Child, M.F. 2016. A conservation assessment of Panthera pardus. In: Child, M.F., Roxburgh, L., Do Linh San, E., Raimondo, D. & Davies-Mostert H.–T. (eds). The Red List of Mammals of South Africa,

19 | P a g e

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Swaziland and Lesotho. South African National Biodiversity Institute and Endangered Wildlife Trust, South Africa. https://www.ewt.org.za/reddata/pdf/RLA_Acinonyx%20jubatus_VU.pdf

Sweanor, L.L., Logan, K.A. & Hornocker, M.G. 2000. Cougar dispersal patterns, Metapopulation dynamics, and conservation. Conservation Biology 14: 798–808.

Treves, A., Krofel, M. and McManus, J., 2016. Predator control should not be a shot in the dark. Frontiers in Ecology and the Environment, 14(7), pp.380-388.

Van der Merwe, V., Marnewick, K., Bissett, C., Groom, R., Mills, M.G.L. & Durant, S.M. 2016. A conservation assessment of Acinonyx jubatus. In: Child, M.F., Roxburgh, L., Do Linh San, E., Raimondo, D. & Davies-Mostert H.–T. (eds). The Red List of Mammals of South Africa, Swaziland and Lesotho. South African National Biodiversity Institute and Endangered Wildlife Trust, South Africa. https://www.ewt.org.za/reddata/pdf/RLA_Acinonyx%20jubatus_VU.pdf

Wagner, K.K., Schmidt, R.H. & Conover, M.R. 1997. Compensation programs for wildlife damage in North America. Wildlife Society Bulletin 25: 312–319.

Woodroffe, R., Thirgood, S., & Rabinowitz, A. 2005. The impact of human–wildlife conflict on natural systems. In R. Woodroffe, S. Thirgood, & A. Rabinowitz (Eds.), People and Wildlife, Conflict or Co- existence? (Conservation Biology, pp. 1-12). Cambridge: Cambridge University Press. doi:10.1017/CBO9780511614774.002

Use of Livestock Guarding Dogs to Reduce Human-Cheetah Conflict. In: Nyhus, P.J., Marker, L., Boast, L.K. and Schmidt-Kuentzel, A., 2017. Cheetahs: Biology and Conservation: Biodiversity of the World: Conservation from Genes to Landscapes. Academic Press

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Sources consulted for the formulation of the submission but not necessarily cited in the text may be included (not a requisite) as a separate list.

Preliminary bibliography:

Alerstam, T., Hedenström, A. and Åkesson, S., 2003. Long-distance migration: evolution and determinants. Oikos 103, 247-260.

Alexander, S., Logan, T. and Paquet, P., 2006. Spatio-temporal co-occurrence of cougars (Felis concolor), wolves (Canis lupus) and their prey during winter: a comparison of two analytical methods. Journal of Biogeography 33, 2001-2012.

Anderson, C.D., Epperson, B.K., Fortin, M.J., Holderegger, R., James, P., Rosenberg, M.S., Scribner, K.T. and Spear, S., 2010, Considering spatial and temporal scale in landscape-genetic studies of gene flow. Molecular Ecology 19, 3565-3575.

Anderson, D.R. and Burnham, K.P., 2002. Avoiding pitfalls when using information-theoretic methods. The Journal of Wildlife Management, 912-918.

Anderson, D.R., 1997. Corridor Use, Feeding Ecology, and Habitat Relationships of Black Bears in a Fragmented Landscape in Louisiana. M.Sc. University of Tennessee, Knoxville, Tennessee.

Athreya, V., Odden, M., Linnell, J.D., Krishnaswamy, J. and Karanth, K.U., 2016. A cat among the dogs: leopard Panthera pardus diet in a human-dominated landscape in western Maharashtra, India. Oryx, 50(1), pp.156-162.

Bailey, T.N., 1993, The African leopard: ecology and behavior of a solitary felid. Columbia University Press.

Balkenhol, N., Waits, L.P. and Dezzani, R.J., 2009, Statistical approaches in landscape genetics: an evaluation of methods for linking landscape and genetic data. Ecography 32, 818-830.

Balme, G.A., Slotow, R. and Hunter, L.T., 2009. Impact of conservation interventions on the dynamics and persistence of a persecuted leopard (Panthera pardus) population. Biological Conservation 142, 2681-2690.

Balme, G., Slotow, R. and Hunter, L., 2010. Edge effects and the impact of non-protected areas in carnivore conservation: leopards in the Phinda–Mkhuze Complex, South Africa. Animal Conservation 13, 315-323.

Balme, G.A. and Hunter, L.T., 2013. Why leopards commit infanticide. Animal behaviour, 86(4), pp.791-799.

Basille, M., Herfindal, I., Santin-Janin, H., Linnell, J.D., Odden, J., Andersen, R., Arild Høgda, K. and Gaillard, J.M., 2009. What shapes Eurasian lynx distribution in human dominated landscapes: selecting prey or avoiding people? Ecography 32, 683-691.

Bates, D., Maechler, M., Bolker, B., Walker, S., Christensen, R.H.B., Singmann, H., Dai, B., Grothendieck, G., Green, P. and Bolker, M.B., 2014. Package ‘lme4’. R foundation for statistical computing, Vienna.

Battin, J., 2004. When good animals love bad habitats: ecological traps and the conservation of animal populations. Conservation Biology 18, 1482-1491.

21 | P a g e

WAPFSA Leopard Trophy Hunting Quota Submission, 2019

Beausoleil, R.A., Koehler, G.M., Maletzke, B.T., Kertson, B.N. and Wielgus, R.B., 2013. Research to regulation: Cougar social behavior as a guide for management. Wildlife Society Bulletin, 37(3), pp.680-688.

Beier, P., 1996, Metapopulation models, tenacious tracking, and cougar conservation. Metapopulations and Wildlife Conservation. Island Press, Covelo, CA, 293-322.

Beier, P., Spencer, W., Baldwin, R.F. and McRae, B., 2011. Toward best practices for developing regional connectivity maps. Conservation Biology 25, 879-892.

Belovsky, G.E., 1987. Extinction models and mammalian persistence, In: Soule, M.E. (Ed.) Viable populations for conservation. Cambridge University Press, Cambridge, U. K., pp. 35-57.

Berggren, Å., Birath, B. and Kindvall, O., 2002. Effect of corridors and habitat edges on dispersal behavior, movement rates, and movement angles in Roesel’s bush-cricket (Metrioptera roeseli). Conservation Biology 16, 1562-1569.

Boitani, L., Corsi, F., De Biase, A., Carranza, I.D., Ravagli, M., Reggiani, G., Sinibaldi and I., Trapanese, P. 1999. A databank for the conservation and management of the African mammals (Roma, Italy, Instituto di Ecolgia Applicata).

Boshoff, A., Cowling, R.M. and Kerley, G.I.H., 2000. The Baviaanskloof Conservation Area: a conservation and tourism development priority. Terrestrial Ecology Research Unit, University of Port Elizabeth.

Bothma, J.d.P., Knight, M., Le Riche, E. and Van Hensbergen, H., 1997. Range size of southern Kalahari leopards. South African Journal of Wildlife Research 27, 94-99.

Bowler, D.E. and Benton, T.G., 2005. Causes and consequences of animal dispersal strategies: relating individual behaviour to spatial dynamics. Biological Reviews 80, 205-225.

Boyce, M.S. and McDonald, L.L., 1999. Relating populations to habitats using resource selection functions. Trends in Ecology & Evolution 14, 268-272.

Boyce, M.S., 2006. Scale for resource selection functions. Diversity and Distributions 12, 269-276.

Boyce, M.S., Vernier, P.R. and Nielsen, S.E., Schmiegelow, F.K., 2002. Evaluating resource selection functions. Ecological modelling 157, 281-300.

Boydston, E.E., Kapheim, K.M., Watts, H.E., Szykman, M. and Holekamp, K.E., 2003. Altered behaviour in spotted hyenas associated with increased human activity. Animal Conservation 6, 207- 219.

Broomhall, L., Mills, M. andDu Toit, J., 2004. Home range and habitat use by cheetahs (Acinonyx jubatus) in the Kruger National Park. Journal of Zoology 261, 119-128.

Bunnefeld, N., Linnell, J.D., Odden, J., Van Duijn, M. and Andersen, R., 2006. Risk taking by Eurasian lynx (Lynx lynx) in a human-dominated landscape: effects of sex and reproductive status. Journal of Zoology 270, 31-39.

Bunnefeld, N., Linnell, J.D., Odden, J., Van Duijn, M. and Andersen, R., 2006. Risk taking by Eurasian lynx (Lynx lynx) in a human-dominated landscape: effects of sex and reproductive status. Journal of Zoology 270, 31-39.

22 | P a g e

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Cantú-Salazar, L. and Gaston, K.J., 2010. Very large protected areas and their contribution to terrestrial biological conservation. Bioscience 60, 808-818.

Carbone, C. and Gittleman, J.L., 2002. A common rule for the scaling of carnivore density. Science 295, 2273-2276.

Carroll, C., Noss, R.F. and Paquet, P.C., 2001. Carnivores as focal species for conservation planning in the Rocky Mountain region. Ecological applications 11, 961-980.

Carroll, C., Noss, R.F.and Paquet, P.C., 2001, Carnivores as focal species for conservation planning in the Rocky Mountain region. Ecological applications 11, 961-980.

Cegelski, C., Waits, L. and Anderson, N., 2003, Assessing population structure and gene flow in Montana wolverines (Gulo gulo) using assignment-based approaches. Molecular Ecology 12, 2907- 2918.

Chetkiewicz, C.-L.B., St. Clair, C.C. and Boyce, M.S., 2006. Corridors for conservation: integrating pattern and process. Annual Review of Ecology, Evolution, and Systematics, 317-342.

Chruszcz, B., Clevenger, A.P., Gunson, K.E. and Gibeau, M.L., 2003. Relationships among grizzly bears, highways, and habitat in the Banff-Bow Valley, Alberta, Canada. Canadian Journal of Zoology 81, 1378-1391.

CIA 2003. The World Factbook – Namibia (http://www.cia.gov/cia/publications/factbook/geos/wa.html).

Clark, T.W., Curlee, A.P. and Reading, R.P., 1996. Crafting effective solutions to the large carnivore conservation problem. Conservation Biology 10, 940-948.

Codling, E.A., Plank, M.J. and Benhamou, S., 2008. Random walk models in biology. Journal of the Royal Society Interface 5, 813-834.

Colchero, F., Conde, D.A., Manterola, C., Chávez, C., Rivera, A. and Ceballos, G., 2011. Jaguars on the move: modeling movement to mitigate fragmentation from road expansion in the Mayan Forest. Animal Conservation 14, 158-166.

Conde, D.A., Colchero, F., Zarza, H., Christensen, N.L., Sexton, J.O., Manterola, C., Chávez, C., Rivera, A., Azuara, D. and Ceballos, G., 2010. Sex matters: Modeling male and female habitat differences for jaguar conservation. Biological Conservation 143, 1980-1988.

Creel, S. and Creel, N.M., 1995. Communal hunting and pack size in African wild dogs, Lycaon pictus. Animal Behaviour 50, 1325-1339.

Creel, S., Spong, G. and Creel, N., 2001. Interspecific competition and the population biology of extinction-prone carnivores. Conservation Biology Series. Cambridge, pp 35 – 60

Crook, J.H., 1970. Social organization and the environment: aspects of contemporary social ethology. Animal Behaviour, 18, pp.197-209.

Cushman, S.A., McRae, B., Adriansen, F., Beier, P., Shirley, M. and Zeller, K., 2013. Biological corridors and connectivity. Key topics in conservation biology 2, 384-404.

Devens, C., Tshabalala, T., McManus, J.S. and Smuts., B. The use of spatially explicit capture recapture techniques and GPS data to estiamte leoaprd (Panthera pardus) density in the Eastern and Western Cape, South Africa. Submitted to African Journal of Ecology.

23 | P a g e

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Dudley, N., 2008. Guidelines for applying protected area management categories. IUCN. Dudley, N., 2008. Guidelines for applying protected area management categories. IUCN.

Dutta, T., Sharma, S., Maldonado, J.E., Wood, T.C., Panwar, H. and Seidensticker, J., 2013, Fine-scale population genetic structure in a wide-ranging carnivore, the leopard (Panthera pardus fusca) in central India. Diversity and Distributions 19, 760-771.

Edelhoff, H., Signer, J. and Balkenhol, N., 2016. Path segmentation for beginners: an overview of current methods for detecting changes in animal movement patterns. Movement Ecology 4, 21.

Elbroch, L.M. and Lendrum, P.E., Quigley, H. and Caragiulo, A., 2016. Spatial overlap in a solitary carnivore: support for the land tenure, kinship or resource dispersion hypotheses? Journal of Animal Ecology, 85(2), pp.487-496.

Elfström, M., Zedrosser, A., Støen, O.G. and Swenson, J.E., 2014. Ultimate and proximate mechanisms underlying the occurrence of bears close to human settlements: review and management implications. Mammal Review, 44(1), pp.5-18.

Elliot, N.B., Cushman, S.A., Loveridge, A.J., Mtare, G. and Macdonald, D.W., 2014. Movements vary according to dispersal stage, group size, and rainfall: the case of the African lion. Ecology 95, 2860- 2869.

Elmhagen, B. and Rushton, S.P., 2007. Trophic control of mesopredators in terrestrial ecosystems: top-down or bottom-up? Ecology letters 10, 197-206.

Epps, C.W., Palsbøll, P.J., Wehausen, J.D., Roderick, G.K., Ramey, R.R. and McCullough, D.R., 2005, Highways block gene flow and cause a rapid decline in genetic diversity of desert bighorn sheep. Ecology letters 8, 1029-1038.

Epps, C.W., Palsbøll, P.J., Wehausen, J.D., Roderick, G.K., Ramey, R.R. and McCullough, D.R., 2005. Highways block gene flow and cause a rapid decline in genetic diversity of desert bighorn sheep. Ecology letters 8, 1029-1038.

Ernest, H.B., Boyce, W.M., Bleich, V.C., May, B., Stiver, S.J. and Torres, S.G., 2003, Genetic structure of mountain lion (Puma concolor) populations in California. Conservation Genetics 4, 353-366.

Estes, J.A., Terborgh, J., Brashares, J.S., Power, M.E., Berger, J., Bond, W.J., Carpenter, S.R., Essington, T.E., Holt, R.D. and Jackson, J.B., 2011. Trophic downgrading of planet Earth. Science 333, 301-306.

Fahrig, L., 2007. Non-optimal animal movement in human-altered landscapes. Functional Ecology 21, 1003-1015.

Fattebert, J., Robinson, H.S., Balme, G., Slotow, R. and Hunter, L., 2015. Structural habitat predicts functional dispersal habitat of a large carnivore: how leopards change spots. Ecological applications 25, 1911-1921.

Fernández, N., Delibes, M. and Palomares, F., 2006. Landscape evaluation in conservation: molecular sampling and habitat modeling for the Iberian lynx. Ecological applications 16, 1037-1049.

Ferreras, P., 2001. Landscape structure and asymmetrical inter-patch connectivity in a metapopulation of the endangered Iberian lynx. Biological Conservation 100, 125-136.

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WAPFSA Leopard Trophy Hunting Quota Submission, 2019

Ferreras, P., Delibes, M., Palomares, F., Fedriani, J.M., Calzada, J. and Revilla, E., 2004. Proximate and ultimate causes of dispersal in the Iberian lynx Lynx pardinus. Behavioral Ecology 15, 31-40.

Firle, S., Bommarco, R., Ekbom, B. and Natiello, M., 1998. The influence of movement and resting behavior on the range of three carabid beetles. Ecology 79, 2113-2122.

Forester, J.D., Ives, A.R., Turner, M.G., Anderson, D.P., Fortin, D., Beyer, H.L., Smith, D.W. and Boyce, M.S., 2007. State–space models link elk movement patterns to landscape characteristics in Yellowstone National Park. Ecological Monographs 77, 285-299.

Forman, R.T. and Alexander, L.E., 1998. Roads and their major ecological effects. Annual review of ecology and systematics 29, 207-231.

Frair, J.L., Merrill, E.H., Beyer, H.L. and Morales, J.M., 2008. Thresholds in landscape connectivity and mortality risks in response to growing road networks. Journal of Applied Ecology 45, 1504-1513.

Franke, A., Caelli, T. and Hudson, R.J., 2004. Analysis of movements and behavior of caribou (Rangifer tarandus) using hidden Markov models. Ecological modelling 173, 259-270.

Frankham, R., 2005, Genetics and extinction. Biological Conservation 126, 131-140.

Gaines, W.L., Lyons, A.L., Lehmkuhl, J.F. and Raedeke, K.J., 2005. Landscape evaluation of female black bear habitat effectiveness and capability in the North Cascades, Washington. Biological Conservation 125, 411-425.

Garamszegi, L.Z., 2011. Information-theoretic approaches to statistical analysis in behavioural ecology: an introduction. Behavioral Ecology and Sociobiology 65, 1-11.

Gavashelishvili, A. and Lukarevskiy, V., 2008, Modelling the habitat requirements of leopard Panthera pardus in west and central Asia. Journal of Applied Ecology 45, 579-588.

Geffen, E., Hefner, R., Macdonald, D.W. and Ucko, M., 1992. Habitat selection and home range in the Blanford’s fox, Vulpes cana: compatibility with the resource dispersion hypothesis. Oecologia 91, 75- 81.

Gillies, C.S., Hebblewhite, M., Nielsen, S.E., Krawchuk, M.A., Aldridge, C.L., Frair, J.L., Saher, D.J., Stevens, C.E. and Jerde, C.L., 2006. Application of random effects to the study of resource selection by animals. Journal of Animal Ecology 75, 887-898.

Ginsberg, J.R. and Milner-Gulland, E., 1994, Sex-Biased Harvesting and Population Dynamics in Ungulates: Implications for Conservation and Sustainable Use. Conservation Biology 8, 157-166.

Gittleman, J.L., 2013, Carnivore behavior, ecology, and evolution. Springer Science & Business Media.

Gittleman, J.L., Funk, S.M., Macdonald, D.W. and Wayne, R.K. 2001. Carnivore Conservation. In Conservation Biology series (Cambridge, U. K., Cambridge University Press).

Graves, T.A., Farley, S., Goldstein, M.I. and Servheen, C., 2007. Identification of functional corridors with movement characteristics of brown bears on the Kenai Peninsula, Alaska. Landscape Ecology 22, 765-772.

Guillot, G., Mortier and F., Estoup, A., 2005, GENELAND: a computer package for landscape genetics. Molecular ecology resources 5, 712-715.

25 | P a g e

WAPFSA Leopard Trophy Hunting Quota Submission, 2019

Hale, M.L., Burg, T.M. and Steeves, T.E., 2012, Sampling for microsatellite-based population genetic studies: 25 to 30 individuals per population is enough to accurately estimate allele frequencies. PloS one 7, e45170.

Hamilton, W.D., 1964. The genetical evolution of social behaviour. II. Journal of theoretical biology 7, 17-52.

Hanski, I. and Gilpin, M., 1991, Metapopulation dynamics: brief history and conceptual domain. Biological journal of the Linnean Society 42, 3-16.

Hanski, I., 1998. Metapopulation dynamics. Nature 396, 41-49.

Harrison, S. and Taylor, A.D., 1997, Empirical evidence for metapopulation dynamics. Metapopulation biology: ecology, genetics, and evolution, 27-42.

Harrison, S., 1991, Local extinction in a metapopulation context: an empirical evaluation. Biological journal of the Linnean Society 42, 73-88.

Hebblewhite, M. and Merrill, E., 2008. Modelling wildlife–human relationships for social species with mixed-effects resource selection models. Journal of Applied Ecology 45, 834-844.

Hess, G.R. and Fischer, R.A., 2001. Communicating clearly about conservation corridors. Landscape and urban planning 55, 195-208.

Hirzel, A.H. and Le Lay, G., 2008. Habitat suitability modelling and niche theory. Journal of Applied Ecology 45, 1372-1381.

Hoffmann, A.A. and Willi, Y., 2008. Detecting genetic responses to environmental change. Nature Reviews Genetics 9, 421-432.

Holderegger, R. and Di Giulio, M., 2010, The genetic effects of roads: a review of empirical evidence. Basic and Applied Ecology 11, 522-531.

Holyoak, M., Casagrandi, R., Nathan, R., Revilla, E. and Spiegel, O., 2008. Trends and missing parts in the study of movement ecology. Proceedings of the National Academy of Sciences 105, 19060- 19065.

Hopcraft, J.G.C., Morales, J., Beyer, H., Borner, M., Mwangomo, E., Sinclair, A., Olff, H. and Haydon, D.T., 2014. Competition, predation, and migration: individual choice patterns of Serengeti migrants captured by hierarchical models. Ecological Monographs 84, 355-372.

Hornocker, M.G., 1969. Winter territoriality in mountain lions. The Journal of wildlife management, 457-464.

Hunter, M.L., 1999. Maintaining biodiversity in forest ecosystems. Cambridge Univ Pr.

Jacobson AP, Gerngross P, Lemeris Jr. JR, Schoonover RF, Anco C, Breitenmoser-Würsten C, Durant SM, Farhadinia MS, Henschel P, Kamler JF, Laguardia A, Rostro-García S, Stein AB, Dollar L. 2016. Leopard (Panthera pardus) status, distribution, and the research efforts across its range. PeerJ 4:e1974 https://doi.org/10.7717/peerj.1974

Johnson, C.J., Nielsen, S.E., Merrill, E.H., McDonald, T.L. and Boyce, M.S., 2006. Resource selection functions based on use-availability data: theoretical motivation and evaluation methods. Journal of wildlife Management 70, 347-357.

26 | P a g e

WAPFSA Leopard Trophy Hunting Quota Submission, 2019

Johnson, D.H., 1980. The comparison of usage and availability measurements for evaluating resource preference. Ecology 61, 65-71.

Kaartinen, S., Luoto, M. and Kojola, I., 2009. Carnivore-livestock conflicts: determinants of wolf (Canis lupus) depredation on sheep farms in Finland. Biodiversity and Conservation 18, 3503.

Karanth, K.U., Nichols, J.D., Kumar, N.S., Link, W.A. and Hines, J.E., 2004. Tigers and their prey: predicting carnivore densities from prey abundance. Proceedings of the National Academy of Sciences of the United States of America 101, 4854-4858.

Keller, L.F. and Waller, D.M., 2002, Inbreeding effects in wild populations. Trends in Ecology & Evolution 17, 230-241.

Kerk, M., Onorato, D.P., Criffield, M.A., Bolker, B.M., Augustine, B.C., McKinley, S.A. and Oli, M.K., 2015. Hidden semi-Markov models reveal multiphasic movement of the endangered Florida panther. Journal of Animal Ecology 84, 576-585.

Kerley, L.L., Goodrich, J.M., Miquelle, D.G., Smirnov, E.N., Quigley, H.B. and Hornocker, M.G., 2002. Effects of roads and human disturbance on Amur tigers. Conservation Biology 16, 97-108.

Kimura, M. and Ohta, T., 1971, Theoretical aspects of population genetics, Vol 4. Princeton University Press.

Klar, N., Fernández, N., Kramer-Schadt, S., Herrmann, M., Trinzen, M., Büttner, I. and Niemitz, C., 2008. Habitat selection models for European wildcat conservation. Biological conservation, 141(1), pp.308-319.

Knopff, A.A., Knopff, K.H., Boyce, M.S. and Clair, C.C.S., 2014. Flexible habitat selection by cougars in response to anthropogenic development. Biological Conservation 178, 136-145.

Koper, N. and Manseau, M., 2009. Generalized estimating equations and generalized linear mixed- effects models for modelling resource selection. Journal of Applied Ecology 46, 590-599.

Krebs, C.J., Boutin, S., Boonstra, R., Sinclair, A.R.E., Smith, J.N.M., Dale, M.R.T., Martin, K. and Turkington, R., 1995. Impact of food and predation on the snowshoe hare cycle. Science 269, 1112- 1118.

Lande, R. and Barrowclough, G.F., 1987, Effective population size, genetic variation, and their use in population management. Viable populations for conservation 87, 124.

Langrock, R., King, R., Matthiopoulos, J., Thomas, L., Fortin, D. and Morales, J.M., 2012. Flexible and practical modeling of animal telemetry data: hidden Markov models and extensions. Ecology 93, 2336-2342.

Levins, R., 1970, Extinction. Some mathematical problems in biology. American Mathematical Society, Providence, Rhode Island, 77-107.

Levins, R., 1968. Evolution in changing environments: some theoretical explorations. Princeton University Press.

Linkie, M., Chapron, G., Martyr, D.J., Holden, J. and Leader-Williams, N, 2006. Assessing the viability of tiger subpopulations in a fragmented landscape. Journal of Applied Ecology, 43(3), pp.576-586.

Logan, K.A. and Sweanor, L.L., 2001. Desert Puma: Evolutionary Ecology and Conservation of an Enduring Carnivore. Island Press, Washington D.C.

27 | P a g e

WAPFSA Leopard Trophy Hunting Quota Submission, 2019

Logan, K.A., Sweanor, L.L., Hornocker, M. and Negri, S., 2009. Behavior and social organization of a solitary carnivore. Cougar, ecology and conservation (M. Hornocker and S. Negri, eds.). University of Chicago Press, Chicago, Illinois, 105-117.

Loxterman, J.L., 2011, Fine scale population genetic structure of pumas in the Intermountain West. Conservation Genetics 12, 1049-1059.

Mace, R.D., Waller, J.S., Manley, T.L., Lyon, L.J. and Zuuring, H., 1996. Relationships among grizzly bears, roads and habitat in the Swan Mountains Montana. Journal of Applied Ecology, 1395-1404.

Manly, B., McDonald, L., Thomas, D., McDonald, T. and Erickson, W., 2002. Resource selection by animals: statistical analysis and design for field studies. Nordrecht, The Netherlands: Kluwer.

Mann, G. 2014. Aspects of the ecology of leopards (Panthera pardus) in the Little Karoo, South Africa. PhD. Thesis. Rhodes University, Grahamstown, South Africa.

Marker, L., 2002. Aspects of cheetah (Acinonyx jubatus) biology, ecology and conservation strategies on Namibian farmlands. D. Phil. University of Oxford, Oxford, U.K.

Marshal, J.P., Bleich, V.C., Krausman, P.R., Reed, M.L. and Andrew, N.G., 2006. Factors affecting habitat use and distribution of desert mule deer in an arid environment. Wildlife Society Bulletin 34, 609-619.

Martin, J., Basille, M., Van Moorter, B., Kindberg, J., Allaine, D. and Swenson, J.E., 2010. Coping with human disturbance: spatial and temporal tactics of the brown bear (Ursus arctos). Canadian Journal of Zoology 88, 875-883.

Martins, Q. 2010. The ecology of the leopard (Panthera pardus) in the Cederberg Mountains. Bristol University, Briston, United Kingdom.

McKellar, A.E., Langrock, R., Walters, J.R. and Kesler, D.C., 2015. Using mixed hidden Markov models to examine behavioral states in a cooperatively breeding bird. Behavioral Ecology 26, 148-157.

McManus, J., Dickman, A., Gaynor, D., Smuts, B. and Macdonald, D., 2014, Dead or alive? Comparing costs and benefits of lethal and non-lethal human–wildlife conflict mitigation on livestock farms. Oryx, 1-9.

McRae, B., Beier, P., Dewald, L., Huynh, L., Keim, P., 2005, Habitat barriers limit gene flow and illuminate historical events in a wide-ranging carnivore, the American puma. Molecular Ecology 14, 1965-1977.

Mech, L.D., 1989. Wolf population survival in an area of high road density. American Midland Naturalist, 387-389.

Mech, L.D., Fritts, S.H. and Wagner, D., 1995. Minnesota wolf dispersal to Wisconsin and Michigan. American Midland Naturalist, 368-370.

Melnick, D.J. and Hoelzer, G.A., 1992, Differences in male and female macaque dispersal lead to contrasting distributions of nuclear and mitochondrial DNA variation. International Journal of Primatology 13, 379-393.

Menotti-Raymond, M. and O’Brien, S.J., 1994, Evolutionary conservation of ten microsatellite loci in four species of Felidae. The Journal of heredity 86, 319-322.

28 | P a g e

WAPFSA Leopard Trophy Hunting Quota Submission, 2019

Michelot, T., Langrock, R. and Patterson, T.A., 2016. moveHMM: An R package for the statistical modelling of animal movement data using hidden Markov models. Methods in Ecology and Evolution.

Miller, J.R., 2015. Mapping attack hotspots to mitigate human–carnivore conflict: approaches and applications of spatial predation risk modeling. Biodiversity and Conservation 24, 2887-2911.

Mills, L.S. and Allendorf, F.W., 1996, The one-migrant-per-generation rule in conservation and management. Conservation Biology 10, 1509-1518.

Mills, M.G.L., Freitag, S. and van Jaarsveld, A.S., 2001. Geographic priorities for carnivore conservation in Africa, In: Gittleman, J.L., Funk, S.M., Macdonald, D., Wayne, R.K. (Eds.) Carnivore Conservation. Cambridge University Press, Cambridge, U. K., pp. 467-483.

Miththapala, S., Seidensticker, J. and O’brien, S.J., 1996, Phylogeographic subspecies recognition in leopards (Panthera pardus): molecular genetic variation. Conservation Biology 10, 1115-1132.

Moilanen, A. and Nieminen, M., 2002. Simple connectivity measures in spatial ecology. Ecology, 83(4), pp.1131-1145.

Mondol, S., Navya, R., Athreya, V., Sunagar, K., Selvaraj, V.M. and Ramakrishnan, U., 2009, A panel of microsatellites to individually identify leopards and its application to leopard monitoring in human dominated landscapes. Bmc Genetics 10, 79.

Morales, J.M. and Ellner, S.P., 2002. Scaling up animal movements in heterogeneous landscapes: the importance of behavior. Ecology 83, 2240-2247.

Morris, D.W., 2003. Toward an ecological synthesis: a case for habitat selection. Oecologia 136, 1-13.

Morrison, J.C., Sechrest, W., Dinerstein, E., Wilcove, D.S. and Lamoreux, J.F., 2007. Persistence of large mammal faunas as indicators of global human impacts. Journal of Mammalogy 88, 1363-1380.

Morrison, M.L., 2001. A proposed research emphasis to overcome the limits of wildlife-habitat relationship studies. The Journal of wildlife management, 613-623.

Mucina, L. and Rutherford, M.C., 2006. The vegetation of South Africa, Lesotho and Swaziland. South African National Biodiversity Institute.

Mysterud, A. and Ims, R.A., 1998. Functional responses in habitat use: availability influences relative use in trade-off situations. Ecology 79, 1435-1441.

Nathan, R., Getz, W.M., Revilla, E., Holyoak, M., Kadmon, R., Saltz, D. and Smouse, P.E., 2008. A movement ecology paradigm for unifying organismal movement research. Proceedings of the National Academy of Sciences 105, 19052-19059.

Nathan, R., Getz, W.M., Revilla, E., Holyoak, M., Kadmon, R., Saltz, D. and Smouse, P.E., 2008. A movement ecology paradigm for unifying organismal movement research. Proceedings of the National Academy of Sciences 105, 19052-19059.

Nee, S. and May, R.M., 1992. Dynamics of metapopulations: habitat destruction and competitive coexistence. Journal of Animal Ecology 61, 37-40.

Northrup, J., Stenhouse, G. and Boyce, M., 2012, Agricultural lands as ecological traps for grizzly bears. Animal Conservation 15, 369-377.

29 | P a g e

WAPFSA Leopard Trophy Hunting Quota Submission, 2019

Noss, R.F., Carroll, C., Vance-Borland, K. and Wuerthner, G., 2002, A multicriteria assessment of the irreplaceability and vulnerability of sites in the Greater Yellowstone Ecosystem. Conservation Biology 16, 895-908.

O’Brien, S.J. and Johnson, W.E., 2005, Big cat genomics. Annu. Rev. Genomics Hum. Genet. 6, 407- 429.

Olea, P.P., Mateo-Tomás, P. and De Frutos, Á., 2010. Estimating and modelling bias of the hierarchical partitioning public-domain software: implications in environmental management and conservation. PloS one 5, e11698.

Packer, C. 1986. The ecology of sociality in felids. Ecological Aspects of Social Evolution (eds D.E. Rubenstein and R.W. Wrangham), pp 429 – 451. Princeton University Press, Princeton, USA.

Palomares, F., Delibes, M., Ferreras, P., Fedriani, J.M., Calzada, J. and Revilla, E., 2000. Iberian lynx in a fragmented landscape: predispersal, dispersal, and postdispersal habitats. Conservation Biology, 14(3), pp.809-818.

Parejo, D., Danchin, E. and Avilés, J.M., 2005. The heterospecific habitat copying hypothesis: can competitors indicate habitat quality? Behavioral Ecology 16, 96-105.

Patterson, T.A., Basson, M., Bravington and M.V., Gunn, J.S., 2009. Classifying movement behaviour in relation to environmental conditions using hidden Markov models. Journal of Animal Ecology 78, 1113-1123.

Patterson, T.A., Thomas, L., Wilcox, C., Ovaskainen, O. and Matthiopoulos, J., 2008. State–space models of individual animal movement. Trends in Ecology & Evolution 23, 87-94.

Peakall, R. and Smouse, P.E., 2006, GENALEX 6: genetic analysis in Excel. Population genetic software for teaching and research. Molecular ecology notes 6, 288-295.

Pilot, M., Jedrzejewski, W., Branicki, W., Sidorovich, V.E., Jedrzejewska, B., Stachura, K. and Funk, S.M., 2006, Ecological factors influence population genetic structure of European grey wolves. Molecular Ecology 15, 4533-4553.

Pohle, J., Langrock, R., Van Beest, F. and Schmidt, N.M., 2017. Selecting the Number of States in Hidden Markov Models – Pitfalls, Practical Challenges and Pragmatic Solutions. arXiv preprint arXiv:1701.08673.

Popov, V., Langrock, R., DeRuiter, S.L. and Visser, F., 2017. An analysis of pilot whale vocalization activity using hidden Markov models. The Journal of the Acoustical Society of America 141, 159-171.

Powell, R.A., Zimmerman, J.W., Seaman, D.E. and Gilliam, J.F., 1996, Demographic analyses of a hunted black bear population with access to a refuge. Conservation Biology, 224-234.

Pritchard, J.K., Stephens and M., Donnelly, P., 2000, Inference of population structure using multilocus genotype data. Genetics 155, 945-959.

Pulliam, H.R. and Danielson, B.J., 1991. Sources, sinks, and habitat selection: a landscape perspective on population dynamics. American naturalist, S50-S66.

Purvis, A., Mace, G.M. and Gittleman, J.L., 2001. Past and future carnivore extinctions: a phylogenetic perspective, In: Gittleman, J.L., Funk, S.M., Macdonald, D., Wayne, R.K. (Eds.) Carnivore Conservation. Cambridge University Press, Cambridge, U. K., pp. 11-34.

30 | P a g e

WAPFSA Leopard Trophy Hunting Quota Submission, 2019

R Development Core Team, 2014. R: A language and environment for statistical computing (Vienna, Austria).

Rabinowitz, A. and Winter, S., 2006. Connecting the Dots: Paseo Tigre-the Path of the Jaguar-is an ambitious project to maintain corridors connecting habitat throughout Tropical America. WILDLIFE CONSERVATION 109, 24.

Ray, J.C., Hunter, L. and Zigouris, J., 2005. Setting conservation and research priorities for larger African carnivores, Vol 24. Wildlife Conservation Society New York.

Raymond, M. and Rousset, F., 1995, GENEPOP (version 1.2): population genetics software for exact tests and ecumenicism. Journal of heredity 86, 248-249.

Rettie, W.J. and Messier, F., 2000. Hierarchical habitat selection by woodland caribou: its relationship to limiting factors. Ecography, 466-478.

Revilla, E. and Wiegand, T., 2008. Individual movement behavior, matrix heterogeneity, and the dynamics of spatially structured populations. Proceedings of the National Academy of Sciences 105, 19120-19125.

Revilla, E. and Wiegand, T., 2008. Individual movement behavior, matrix heterogeneity, and the dynamics of spatially structured populations. Proceedings of the National Academy of Sciences 105, 19120-19125.

Revilla, E., Wiegand, T., Palomares, F., Ferreras, P. and Delibes, M., 2004. Effects of matrix heterogeneity on animal dispersal: from individual behavior to metapopulation-level parameters. The American Naturalist 164, E130-E153.

Ripple, W.J., Estes, J.A., Beschta, R.L., Wilmers, C.C., Ritchie, E.G., Hebblewhite, M., Berger, J., Elmhagen, B., Letnic, M. and Nelson, M.P., 2014. Status and ecological effects of the world’s largest carnivores. Science 343, 1241484.

Ripple, W.J., Estes, J.A., Beschta, R.L., Wilmers, C.C., Ritchie, E.G., Hebblewhite, M., Berger, J., Elmhagen, B., Letnic, M. and Nelson, M.P., 2014. Status and ecological effects of the world’s largest carnivores. Science 343, 1241484.

Ripple, W.J., Estes, J.A., Beschta, R.L., Wilmers, C.C., Ritchie, E.G., Hebblewhite, M., Berger, J., Elmhagen, B., Letnic, M. and Nelson, M.P., 2014. Status and ecological effects of the world’s largest carnivores. Science 343, 1241484.

Robertson, B.A. and Hutto, R.L., 2006. A framework for understanding ecological traps and an evaluation of existing evidence. Ecology 87, 1075-1085.

Robin, X., Turck, N., Hainard, A., Tiberti, N., Lisacek, F., Sanchez, J.-C. and Müller, M., 2011. pROC: an open-source package for R and S+ to analyze and compare ROC curves. BMC bioinformatics 12, 77.

Rodgers, A.R., Carr, A., Beyer, H., Smith, L. and Kie, J. 2007. HRT: home range tools for ArcGIS (Version).

Roever, C., Beyer, H., Chase, M. and Aarde, R., 2014. The pitfalls of ignoring behaviour when quantifying habitat selection. Diversity and Distributions 20, 322-333.

Roever, C.L., Boyce, M.S. and Stenhouse, G.B., 2010. Grizzly bear movements relative to roads: application of step selection functions. Ecography 33, 1113-1122.

31 | P a g e

WAPFSA Leopard Trophy Hunting Quota Submission, 2019

Romain-Bondi, K.A., Wielgus, R.B., Waits, L., Kasworm, W.F., Austin, M. and Wakkinen, W., 2004. Density and population size estimates for North Cascade grizzly bears using DNA hair-sampling techniques. Biological Conservation 117, 417-428.

Rutherford, M.C., Mucina, L. and Powrie, L.W., 2006. Biomes and bioregions of southern Africa. The vegetation of South Africa, Lesotho and Swaziland. Strelitzia 19, 30-51.

Sala, E., 2006. Top predators provide insurance against climate change. Trends in Ecology & Evolution 21, 479-480.

Sala, O.E., Chapin, F.S., Armesto, J.J., Berlow, E., Bloomfield, J., Dirzo, R., Huber-Sanwald, E., Huenneke, L.F., Jackson, R.B. and Kinzig, A., 2000. Global biodiversity scenarios for the year 2100. Science 287, 1770-1774.

Sandell, M., 1989. The mating tactics and spacing patterns of solitary carnivores. In Carnivore behavior, ecology, and evolution. pp. 164-182. Springer, Boston, MA.

Schaffer, M.E., 1988, Evolutionarily stable strategies for a finite population and a variable contest size. Journal of theoretical biology 132, 469-478.

Schlaepfer, M.A., Runge, M.C. and Sherman, P.W., 2002. Ecological and evolutionary traps. Trends in Ecology & Evolution 17, 474-480.

Schmidt, K., Jedrzejewski, W. and Okarma, H., 1997. Spatial organization and social relations in the Eurasian lynx population in Bialowieza Primeval Forest, Poland. Acta Theriologica 42, 289-312.

Seidensticker, J.C., Hornocker, M.G., Wiles, W.V. and Messick, J.P., 1973. Mountain lion social organization in the Idaho Primitive Area. Wildlife Monographs, 3-60.

Serrouya, R., McLellan, B.N., Boutin, S., Seip, D.R. and Nielsen, S.E., 2011. Developing a population target for an overabundant ungulate for ecosystem restoration. Journal of Applied Ecology 48, 935- 942.

Seutin, G., White, B.N. and Boag, P.T., 1991, Preservation of avian blood and tissue samples for DNA analyses. Canadian Journal of Zoology 69, 82-90.

Sinclair, E.A., Swenson, E.L., Wolfe, M.L., Choate, D.C., Bates, B. and Crandall, K.A., 2001, Gene flow estimates in Utah’s cougars imply management beyond Utah. Animal Conservation 4, 257-264.

Singleton, P.H. and Lehmkuhl, J.F., 2001. Using weighted distance and least-cost corridor analysis to evaluate regional-scale large carnivore habitat connectivity in Washington. UC Davis: Road Ecology Center. Retrieved from: http://www. escholarship. org/uc/item/526536d6.

Skead, C.J., Boshoff, A., Kerley, G.I.H. and Lloyd, P., 2007. Historical incidence of the larger land mammals in the broader Eastern Cape, Vol 13. Centre for African Conservation Ecology, Nelson Mandela Metropolitan University Port Elizabeth.

Sork, V.L., Nason, J., Campbell, D.R. and Fernandez, J.F., 1999, Landscape approaches to historical and contemporary gene flow in plants. Trends in Ecology and Evolution 14, 219-224.

Soulé, M.E. and Sanjayan, M., 1998. Ecology: Conservation Targets: Do They Help? Science 279, 2060-2061.

Spong, G., Johansson, M., Björklund, M., 2000, High genetic variation in leopards indicates large and long-term stable effective population size. Molecular Ecology 9, 1773-1782.

32 | P a g e

WAPFSA Leopard Trophy Hunting Quota Submission, 2019

Stein, A.B., Athreya, V., Gerngross, P., Balme, G., Henschel, P., Karanth, U., Miquelle, D., Rostro- Garcia, S., Kamler, J.F., Laguardia, A., Khorozyan, I. and Ghoddousi, A. , 2016. Panthera pardus. (errata version published in 2016) The IUCN Red List of Threatened Species 2016: e.T15954A102421779. Downloaded on 08 January 2017.

Stenseth, N.C., Ehrich, D., Rueness, E.K., Lingjærde, O.C., Chan, K.-S., Boutin, S., O’Donoghue, M., Robinson, D.A., Viljugrein, H. and Jakobsen, K.S., 2004, The effect of climatic forcing on population synchrony and genetic structuring of the Canadian lynx. Proceedings of the National Academy of Sciences of the United States of America 101, 6056-6061.

Stephens, D.W. and Krebs, J.R., 1986. Foraging theory. Princeton University Press.

Stoner, D.C., Wolfe, M.L., Rieth, W.R., Bunnell, K.D., Durham, S.L. and Stoner, L.L., 2013. De facto refugia, ecological traps and the biogeography of anthropogenic cougar mortality in Utah. Diversity and Distributions, 19(9), pp.1114-1124.

Sunquist, M. and Sunquist, F., 2002. Wild Cats of the World. University of Chicago Press, Chicago.

Swanepoel, L.H., Lindsey, P., Somers, M.J., Van Hoven, W. and Dalerum, F., 2013. Extent and fragmentation of suitable leopard habitat in South Africa. Animal Conservation 16, 41-50.

Swanepoel, L.H., Somers, M.J. and Dalerum, F., 2015. Functional responses of retaliatory killing versus recreational sport hunting of leopards in South Africa. PloS one 10, e0125539.

Sweanor, L.L., Logan, K.A. and Hornocker, M.G., 2000, Cougar dispersal patterns, metapopulation dynamics, and conservation. Conservation Biology 14, 798-808.

Tensen, L., Roelofs, D. and Swanepoel, L.H., 2011. A note on the population structure of leopards (Panthera pardus) in South Africa. South African Journal of Wildlife Research 44, 193-197.

Terborgh, J., 1992. Maintenance of diversity in tropical forests. Biotropica 24, 283-292.

Terborgh, J., Estes, J.A., Paquet, P., Ralls, K., Boyd-Heger, D., Miller, B.J. and Noss, R.F., 1999. The role of top carnivores in regulating terrestrial ecosystems, In: Soule, M.E., Terborgh, J. (Eds.) Continental Conservation: Scientific Foundations of Regional Reserve Networks. Island Press, Washington, pp. 39-64.

Teske, P.R., Papadopoulos, I., Barker, N.P. and McQuaid, C.D., 2012, Mitochondrial DNA paradox: sex-specific genetic structure in a marine mussel–despite maternal inheritance and passive dispersal. Bmc Genetics 13, 45.

Thamm, A. and Johnson, M., 2006. The Cape supergroup, In: Johnson, M.R., Anhaeusser, C.R. and Thomas, R.J. (Eds) The Geology of South Africa. Council for Geoscience, 443-459.

Thamm, A.G. and Johnson, M.R. 2006. The Cape Supergroup. In: The geology of South Africa, (eds) M. R. Johnson, C. R. Annhaeusser & R. J. Thomas, pp. 443–461. Geological Society of South Africa, Johannesburg/ Council for Geoscience, Pretoria, Cape Town.

Tilman, D., 1994. Competition and biodiversity in spatially structured habitats. Ecology 75, 2-16.

Towner, A.V., Leos-Barajas, V., Langrock, R., Schick, R.S., Smale, M.J., Kaschke, T., Jewell, O.J. and Papastamatiou, Y.P., 2016. Sex-specific and individual preferences for hunting strategies in white sharks. Functional Ecology.

Treves, A. and Bruskotter, J., 2014. Tolerance for predatory wildlife. Science 344, 476-477.

33 | P a g e

WAPFSA Leopard Trophy Hunting Quota Submission, 2019

Treves, A., Martin, K.A., Wydeven, A.P. and Wiedenhoeft, J.E., 2011. Forecasting environmental hazards and the application of risk maps to predator attacks on livestock. Bioscience 61, 451-458.

Trivers, R.L., 1976. Sexual selection and resource-accruing abilities in Anolis garmani. Evolution, 253- 269.

Turchin, P., 1991. Translating foraging movements in heterogeneous environments into the spatial distribution of foragers. Ecology 72, 1253-1266.

Turchin, P., 1998. Quantitative analysis of movement: measuring and modeling population redistribution in animals and plants, Vol 1. Sinauer Associates Sunderland.

Uphyrkina, O. and Johnson, W.E., Quigley, H., Miquelle, D., Marker, L., Bush, M., O’Brien, S.J., 2001, Phylogenetics, genome diversity and origin of modern leopard, Panthera pardus. Molecular Ecology 10, 2617-2633.

Van Oosterhout, C., Hutchinson, W.F., Wills, D.P. and Shipley, P., 2004, MICRO-CHECKER: software for identifying and correcting genotyping errors in microsatellite data. Molecular ecology notes 4, 535-538.

Venette, R.C., Kriticos, D.J., Magarey, R.D., Koch, F.H., Baker, R.H., Worner, S.P., Raboteaux, N.N.G., McKenney, D.W., Dobesberger, E.J. andYemshanov, D., 2010. Pest risk maps for invasive alien species: a roadmap for improvement. Bioscience 60, 349-362.

Vucetich, J.A. and Waite, T.A., 2000, Is one migrant per generation sufficient for the genetic management of fluctuating populations? Animal Conservation 3, 261-266.

Walker, C.W., Harveson, L.A., Pittman, M.T., Tewes, M.E. and Honeycutt, R.L., 2000, Microsatellite variation in two populations of mountain lions (Puma concolor) in Texas. The Southwestern Naturalist, 196-203.

Weir, B.S. and Hill, W.G., 2002, Estimating F-statistics. Annual review of genetics 36, 721-750.

Wielgus, R. and Bunnell, F., 1994, Dynamics of a small, hunted brown bear Ursus arctos population in southwestern Alberta, Canada. Biological Conservation 67, 161-166.

Wiens, J.A., Chr, N., Van Horne, B. and Ims, R.A., 1993. Ecological mechanisms and landscape ecology. Oikos, 369-380.

Wilmers, C.C. and Getz, W.M., 2005. Gray wolves as climate change buffers in Yellowstone. PLoS Biology 3, e92.

Wilmers, C.C., Wang, Y., Nickel, B., Houghtaling, P., Shakeri, Y., Allen, M.L., Kermish-Wells, J., Yovovich, V. and Williams, T., 2013. Scale dependent behavioral responses to human development by a large predator, the puma. PloS one 8, e60590.

Wilson, E.O. and MacArthur, R.H., 1967. The theory of island biogeography. Princeton, NJ.

Woodroffe, R. and Ginsberg, J., Macdonald, D., 1997. The African Wild Dog: Status Survey and Conservation Action Plan. IUCN/SSC Canid Specialist Group, Gland, Switzerland.

Woodroffe, R. and Ginsberg, J.R., 1998, Edge effects and the extinction of populations inside protected areas. Science 280, 2126-2128.

34 | P a g e

WAPFSA Leopard Trophy Hunting Quota Submission, 2019

Woodroffe, R., 2000. Predators and people: using human densities to interpret declines of large carnivores. Animal Conservation 3, 165-173.

Woodroffe, R., 2001. Strategies for carnivore conservation: Lessons from contemporary extinctions, In: Gittleman, J.L., Wayne, R.K., Macdonald, D.W., Funk, S.M. (Eds.) Carnivore Conservation. Cambridge University Press, Cambridge, pp. 61-92.

Wright, S., 1984, Evolution and the genetics of populations, volume 3: experimental results and evolutionary deductions, Vol 3. University of Chicago press.

Zeller, K.A., McGarigal, K. and Whiteley, A.R., 2012. Estimating landscape resistance to movement: a review. Landscape Ecology 27, 777-797.

Zhang, D.X. and Hewitt, G.M., 2003, Nuclear DNA analyses in genetic studies of populations: practice, problems and prospects. Molecular Ecology 12, 563-584.

Zollner, P.A. and Lima, S.L., 2005. Behavioral tradeoffs when dispersing across a patchy landscape. Oikos 108, 219-230.

Zucchini, W. and MacDonald, I.L., 2009. Hidden Markov models for time series: an introduction using R, Vol 150. CRC press.

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SANBI •Biodiversity for Life
South African National Biodiversity Institute

South African Leopard Monitoring Project

Michele Pfab ScientificCo-ordinator: ScientificAuthority

4 June 2019


Adaptive management framework for trophy hunting

  • Hunting restricted to leopard hunting zones where scientifically robust data on leopard density trends·indicate overall stable (or increasing) populations
  • Leopard population trends monitored~ standardised, rigorous framework~ systematic camera-trap surveys (60-92 digital camera- traps at 30-46 stations; 2-3 km apart; <60 days)
  • Oversight provided by scientific steering committee for leopard monitoring
  • Best-practice guidelines for monitoring leopards in South Africa

Sooth African National Biodiversity Institute


Monitoring sites

  • Identified based on an existing habitat suitability map~State-and province-run protected areas~ Private conservation areas
    ~ Private conservancies & game ranches~ Community reserves
  • Large enough to accurately estimate leopard population density- at least as large as the maximum home range size of the target species(“‘l00km2)
  • Stable land tenure to ensure continuity in the monitoringSANBI···

South Affkan National Biodlvenity Institute


South African National BiodiYer$ity Institute





0 125 250

500 KilometersI


Data analysis

• Leopard population density estimated using maximum-likelihood SCR methods

• Multi state models fitted to monitoring sites wit.h more than two years of monitoring data to determine trends.


Precautionary safeguards
I. Only hunting zones where trends in leopard

density have been monitored for three or more years are considered for the quota;

  1. The areal extent of a hunting zone isdetermined by leopard density – the lowe,r the density, the larger the zone, and thus less leopards can be hunted (only one leopard rr1aybe hunted per zone);
  2. Only males that are seven years or older rnay be hunted, as their removal has the least impact 011a population.

South Afric.in National Biodiversity Institute




Leopard Quota Review: South Africa 1. History and rationale of leopard export quota

The leopard export quota was originally introduced at the fourth meeting of the Conference of the Parties to CITES (Gaborone, 1983) with Resolution Conf. 4.13, and at CoP7 (Lausanne, 1989) an export quota of 50 leopard hunting trophies and skins for personal use was recommended for South Africa. This was subsequently increased to 75 at CoP8 (Kyoto, 1992).

A Population and Habitat Viability Analysis (PHVA) informed the increase in South Africa’s export quota for leopard hunting trophies and skins for personal use at CoP13 (Bangkok, 2004), to 150, as contained in Resolution Conf. 10.14 (Rev. CoP16). Model scenarios tested in the PHVA, ranging from 0 to150 leopard hunted, showed that an increase in the export quota from 75 to 150 did not increase the risk of extinction of the national population within a time period of 100 years, though a reduction in the size of the population from 93% to 64% of the carrying capacity was predicted along with possible local extinctions.

2. Principles

In reviewing the leopard export quota for South Africa, the following principles are deemed to be important:

  1. Management and conservation of leopard in South Africa is underpinned by the sustainable use paradigm.
  2. South Africa has adopted an adaptive management framework for the management and utilization of leopard.
  3. Monitoring is a crucial component of an adaptive management framework and the involvement of stakeholders in monitoring the resource base should be encouraged.
  4. Management and utilization of leopard in South Africa should be underpinned by robust science.
  5. A variety of socio-ecological / socio-economic contexts in range States necessitate different management approaches, tools and interventions.
  6. It is important to take into account the impact of the illegal leopard skin trade on regional leopard populations.
  7. Well-managed sport hunting is an important conservation tool.
  8. Well-managed sport hunting benefits local livelihoods.



3. Status of leopard in South Africa

a. Distribution of leopard within South Africa
Based on maximum entropy models, Swanepoel, et al., (2013) estimated that approximately 20% (248,770 km2) of South Africa is suitable for leopard (Fig. 1). Suitable habitat is fragmented into four general regions: one stretching along the southeast coast, one occurring in the interior of KwaZulu-Natal, one encompassing the Kruger National Park and interior of Limpopo, and one in the northern region where the Kgalagadi Transfrontier National Park is located (Fig. 1). Approximately 32% of the suitable leopard habitat is situated in protected areas.

Figure 1: Suitable leopard habitat in South Africa predicted from a model containing the full set of environmental variables (land cover; NDVI, gazing capacity, elevation, surface ruggedness, distance to nearest river, human density, distance to roads, distance to villages, cattle density, and small ruminant density). (Copied from Swanepoel, et al., 2013).

b. Leopard abundance in South Africa
Leopards are generally considered uncommon in South Africa, however estimates of the size of the national population vary widely from 2,185 to 23,400 leopards (Martin & De Meulenaer, 1988; Friedmann & Traylor‐Holzer, 2005; Swanepoel, et al., 2014b). None of these estimates are based on rigorous population counts at regional scales, and their confidence intervals are so wide as to make them meaningless (e.g. 2,813-11,632 leopards estimated by Swanepoel, et al., 2014b).

Estimated leopard population densities at sites surveyed in Mpumalanga, KwaZulu-Natal, Limpopo and North West Province vary from 0.2 ± 0.0 leopards/100 km2 (Khamab, North West Province), to as high as 12.2 ± 2.4 leopards/100 km2 (Sabi Sands, Mpumalanga) (Mann, et al., 2017). A separate study in the Maputaland Conservation Unit in KwaZulu- Natal estimated leopard density to range between 1.6 ± 0.62/100 km2 in the smallest protected area (Ndumo) to 8.4 ± 1.03/100 km2 in the largest (western shores, St Lucia) (Ramesh, et al., 2017). A density of 10.7 leopard/100 km2 (Chase Grey, et al., 2013) was



recorded in the montane habitat of the western Soutpansberg, though this is likely to be an overestimate since the survey area in this study was smaller than the expected home range size of a single male leopard. Leopard densities across the Eastern and Western Cape have been estimated between 0.95 leopard/100 km2 and 1.11 leopard/100 km2.

c. Leopard population trends in South Africa
Data generated from the South African Leopard Monitoring Project suggest an 8% decline in the national population per year, and significant declines in leopard density have been observed in five out of 18 sites surveyed in 2017 (Mann, et al., 2017). Some stable populations (e.g. Somkhanda and Manyoni) appear to be well below their potential capacities, while other sites such as Barberton and Songimvelo in Mpumalanga, Ophate in KwaZulu-Natal and Dinokeng in Gauteng appear to no longer have functioning leopard populations (Mann, et al., 2017).

Up to the end of 2016, leopard densities at most monitoring sites in KwaZulu-Natal were relatively stable, with the exception of Hluhluwe-iMfolozi Park and St Lucia Eastern Shores, which both showed significant declines from 2015 to 2016, and Phinda Game Reserve, where the population noticeably increased from 2014 to 2016 (Mann, et al., 2017). Leopard density at Hluhluwe-iMfolozi Game Reserve has declined by >70% in five years, from 13 leopards/100 km2 in 2011 to 3 leopards/100 km2 in 2016 (Mann, et al., 2017). Results from the 2017 surveys showed strong declines in leopard density also at Ithala, Tembe and uMkhuze Game Reserves.

Leopard densities at some monitoring sites in Limpopo appear stable, with annual fluctuations within the standard deviation of previous estimates (Mann, et al., 2017). The notable exceptions to this are Zingela and Makalali Game Reserves, where marked declines in leopard density were recorded from 2016 to 2017 (Mann, et al., 2017). A separate study showed a 44% decline in leopard densities in the Soutpansberg Mountains from 2012 (6.55 leopard/100 km2) to 2016 (3.65 leopard/100 km2) (Williams, et al., 2017).

The leopard populations in the Kruger National Park and surrounding private nature reserves, as well as Loskop Dam Nature Reserve in Mpumalanga, remain stable (Mann, et al., 2017). Though density estimate in Timbavati Game Reserve declined markedly in 2016 to 7.3 ± 1.3 leopards/100 km2 from the ~10.5 leopards/100 km2 recorded between 2013 and 2015, and has remained at this level since (Mann, et al., 2017).

The leopard population in Kwande Nature Reserve in the Eastern Cape appears to be stable, while leopard densities in the Gamkaberg, Rooiberg and Swartberg areas of the Little Karoo, Western Cape, increased from 0.6 ± 0.1 leopards/100 km2 in 2012 to 1.1 ± 0.2 in 2017 (Mann, et al., 2017).

Leopard population trends elsewhere in South Africa are unknown.

d. Threats
At present, the illegal killing of leopards for skins and other body parts for traditional ceremonies and medicines is believed to be the major threat facing leopard within South Africa, and more widely across southern Africa (Hunter, et al., 2013). Leopard skins are used in ceremonial wear by a number of cultural and religious groups in KwaZulu-Natal and Swaziland, most notably the Nazareth Baptist ‘Shembe’ Church. Mark-resight surveys undertaken at religious gatherings of the ‘Shembe’ church suggest that there are



approximately between 13,000 and 18,000 illegal leopard skins in circulation among church members.

Other threats to leopard in South Africa include excessive off‐takes (legal and illegal) of putative damage causing animals (DCAs) (Balme, et al., 2009, St John, et al., 2011, Thorn,et al., 2013, Swanepoel, et al., 2014b; Williams, et al., 2017), the unethical radio-collaring of leopards for research and tourism (Balme, et al., 2014), and habitat loss and fragmentation due to the development of urban areas, mines and agriculture (Di Minin, et al., 2013, Swanepoel, et al., 2013, McManus, et al., 2015; Williams, et al., 2017). In the Western Cape, the loss of wilderness areas is resulting in reduced habitat for leopard prey such as hyrax and small antelopes, increasing the likelihood of leopards relying on livestock for food (Martins & Martins, 2006).

e. Illegal off-take
It is estimated that as many as 1500 – 2500 leopards are illegally harvested annually to meet the demand for skins by the Nazareth Baptist ‘Shembe’ Church; The illegal killing of putative DCAs is typically indiscriminate, the scale of which is currently unknown since illegal off-take of leopards is poorly monitored, if at all.

f. Data quality
Until recently, reliable published information on leopard population sizes and trends at a national scale was poor to non-existent. Detailed estimates of abundance are available for only a very small fraction of the species’ range (e.g. Balme, et al. 2010, Chapman & Balme, 2010, Chase-Grey, et al., 2013, Maputla, et al., 2013; Ramesh, et al., 2017; Williams, et al., 2017). However, in 2013, the provinces of KwaZulu-Natal and Limpopo established leopard monitoring programmes wherein systematic camera-trap surveys were conducted in order to reliably track leopard population trends. The South African Leopard Monitoring Project was recently established through the expansion of these monitoring initiatives to five additional provinces, namely the Eastern Cape, Gauteng, Mpumalanga, North West and the Western Cape (Mann, et al., 2017).

4. Adaptive management framework

a. Management tools/plans
Some provinces in South Africa have adopted management plans that address particular aspects of leopard management (e.g. the Eastern and Western Cape have guidelines for the management of DCAs, while KwaZulu-Natal (Balme, et al., 2010b) and Mpumalanga have specific plans that guide the allocation of hunting quotas). A need for a national management plan that provides standardized guidelines to provinces for the management of the species, particularly in relation to trophy hunting, was identified. To this end, norms and standards for the trophy hunting of leopard in South Africa are currently under development, the general approach of which is to: 1) distribute hunting effort evenly across leopard range in South Africa; 2) limit hunting to male leopards of at least seven years of age; and 3) ensure the mandatory submission of hunt return data and trophy photographs following all leopard hunts (both successful and unsuccessful, including DCA leopards killed). An online application and submission system for the management of trophy hunting has been developed and roll out to the provinces is imminent.



Data from the South African Leopard Monitoring Project are being used to inform the adaptive management of leopard in South Africa, and in particular trophy hunting. The adaptive management framework has been designed to test the impact of leopard hunting within hunting zones, so that quotas can be adjusted annually based on trend data. Hunting will be restricted to leopard hunting zones where scientifically robust data on leopard density trends indicate overall stable (or increasing) populations.

b. Legislative tools
The leopard is listed as vulnerable in terms of section 56 of the National Environmental Management: Biodiversity Act 10 of 2004 (NEMBA) and is regulated in accordance with the Threatened or Protected Species Regulations. Various provincial ordinances and acts provide additional legislative protection. Permits are therefore required to undertake a variety of activities in relation to leopard, e.g. hunting, keeping, selling and other forms of direct use. Permits are issued upon a written application, and each application is handled on its merits in accordance with environmental legislation and policies.

A non-detriment finding (NDF) for leopard, a fundamental requirement for the export of the species in accordance with the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), was approved by the Scientific Authority of South Africa and published for public input in December 2015. Since then, the government of South Africa has made concerted efforts to implement measures recommended in the NDF.

c. Harvest restrictions
Until recently though, there were no restrictions on the age, sex or size of leopards that could be killed (Balme, et al., 2012). The norms and standards for the trophy hunting of leopard will however codify a policy adopted in 2015 to limit hunting to male leopards of at least seven years of age.

d. Compliance and enforcement
The number of leopard trophies hunted in South Africa each year is regulated and individuals require a permit to remove a putative DCA. Trophy hunting and legal DCA control is formally managed at the provincial level, and an improvement in compliance is anticipated once the norms and standards for the trophy hunting of leopard are formally gazetted.

Harvest of leopard is however mostly illegal (e.g. illegal retaliatory killing, off-take for skins, incidental snaring) and some of the provinces seems unable to effectively address this.

e. Management of damage causing animals (DCAs)
It has been suggested that the removal of DCAs or the illegal off-take of leopards significantly exceeds off-take from trophy hunting (St John, et al., 2011, Thorn, et al., 2013). However, according to permits issued, a minimum of 46 leopards were killed as DCAs in seven provinces between 2012 and 2016. In some provinces, progress has been made in reducing the number of leopards killed as DCAs. For example, in the Cederberg area of the Western Cape, the recorded removals of problem leopards have declined significantly in recent years, from 7‐8.6 per year during 1950 – 2003, to 0.8 per year from 2004 ‐ 2006 (Martins & Martins, 2006). Similarly, the numbers of problem leopards killed in northern KwaZulu-Natal declined significantly after 2005 (Balme, et al., 2009). In the case of the Western Cape, reduced off‐take appears to have been due to educational efforts by leopard researchers and the development of alternative livestock protection techniques (Martins & Martins, 2006), whereas in northern KwaZulu-Natal it was probably due to the



implementation of a revised protocol for issuing problem leopard permits (Balme, et al., 2009). An increase in the number of DCA applications in Limpopo has been observed in association with the number of game ranches breeding high-value species, and with the use of predator-proof fencing (Pitman, et al., 2016). Leopards were recorded as the most common putative problem animal (68%) in Limpopo between 2003 and 2012, followed by elephant (20%), lion (4%), brown hyena (3%), black-backed jackal (2%), caracal (2%), cheetah (0.5%) and spotted hyena (0.5%) (Pitman, et al., 2016).

Details of leopards killed as DCAs will in future be captured in the online application and submission system that has been developed for the management of leopard trophy hunting, thereby improving monitoring of DCA off-take.

f. Sustainable off-take quotas
The system of allocating leopard hunting quotas in South Africa has been completely overhauled since the South African Leopard Monitoring Project recorded declines in some leopard populations in KwaZulu-Natal and Limpopo. Recent research also suggested that hunting quotas in Limpopo, which accounted for >60% of leopards trophies hunted in South Africa, were unsustainable, particularly if the removal of putative DCAs was taken into account (Pitman, et al., 2015). Anecdotal information from North West similarly suggested that quotas were too high (Power, 2014). A further concern was the clumping of trophy hunts, leading to increased pressure on specific populations.

Leopard hunting quotas are now adjusted annually based on population trend data generated by the South African Leopard Monitoring Project. Hunting will be excluded from any areas where leopard populations are in decline, and hunting will not be allowed in areas where scientifically robust data on leopard population trends are absent. Hunting zones eligible for a quota are thus those where scientifically robust population trend data indicate increasing or stable leopard populations (i.e. no statistically significant difference in observed leopard density over time). In 2016 and 2017, the leopard hunting quota was set at zero to allow time for the recovery of declining populations and improved management of trophy hunting, while for 2018, the Scientific Authority recommended a quota of seven male leopards of >7 years of age (five in Limpopo and two in KwaZulu-Natal) (Fig 2.). Research has shown that since fewer males are required to maintain the same levels of reproduction, leopards are resilient to disturbance if the prime reproductive female life-stage remains intact (Crookes, et al., 1998, Gaona, et al., 1998), while population models show that hunting of leopard can be sustainable when only male leopard 7 years and older are hunted (Packer, et al., 2011).



Figure 2: Eligible leopard hunting zones recommended for 2018.



5. Monitoring methods

The South African Leopard Monitoring Project provides for a standardised, rigorous framework employing systematic camera-trap surveys for the monitoring of leopard population trends throughout South Africa. In 2017, camera-trap surveys coupled with spatial capture-recapture sampling were completed at 24 sites, which included a mix of state- and province-run protected areas, private conservancies, commercial game ranches and community reserves. The total area covered by camera-trap stations amounted to 4,784 km2and the sampling effort comprised 33,605 camera-trap days. Best practice guidelines for leopard monitoring are currently being developed to encourage the private sector to participate in the project.

An online reporting system for trophy hunting applications and hunt return data (including leopards killed as DCAs) has also been developed and roll out to the provinces is imminent. Hunt return data will be used to determine catch-per-unit-effort and harvest composition as indices of relative abundance.

6. Utilization and monitoring of the export quota

In accordance with Resolution Conf. 10.14 (Rev. CoP16), South Africa is allowed to export 150 leopards annually as hunting trophies and skins for personal use, though between 2005 and 2016 only 73 leopards were exported annually, mostly as hunting trophies (Fig. 3) (CITES Trade Database, UNEP World Conservation Monitoring Centre, Cambridge, UK).



Figure 3: South Africa’s utilization of the export quota for leopard hunting trophies and skins for personal use (150), as reported between 2005 and 2016 (CITES Trade Database, UNEP World Conservation Monitoring Centre, Cambridge, UK).

7. Conservation incentives and benefits

In South Africa, privately-owned land is extremely important for the conservation of leopards, and carnivores in general (Friedmann, et al., 2002; Lindsey, Du Toit & Mills, 2004). However, management of carnivore conflict through translocation and killing of DCAs are limiting carnivore persistence on private land (Lindsey, et al., 2004).

Trophy hunting, when well-managed, can be an important tool for the conservation of species and habitats through the provision of financial incentives, especially when revenues are invested back into conservation and when benefits are shared equitably with local communities (e.g. Lindsey, et al., 2007; Dickson, et al., 2009; UNEP-WCMC, 2013; IUCN, 2016), and it has similarly been argued that trophy hunting can foster tolerance towards leopards. Questionnaire surveys have suggested that the attitudes of landowners to leopards is better than for other predator species which cannot be hunted, such as cheetah and wild dogs, and that game ranchers are more tolerant of leopards than livestock farmers



(Lindsey, et al. 2005), however, in northern KwaZulu-Natal, landowners who hunted the most leopards also removed (legally and illegally) the highest numbers of putative DCAs (Balme, et al., 2009). Similar patterns have been observed more widely (Peebles, et al., 2013; Treves & Bruskotter, 2014). In Limpopo Pitman, et al. (2016) demonstrated that the proportional increase in problem animal control of nuisance wildlife such as leopard has far outweighed the proportional increase in game ranching trends towards more intensive practices, suggesting that intolerance is growing. However, the recent shift in South Africa’s game ranching industry to high-value species and colour variants has likely decreased tolerance towards leopards (and other carnivores) and increased levels of retaliatory killing (Thorn, et al., 2013; Pitman, et al., 2016). More research is required to understand the complex relationship between trophy hunting and tolerance of landowners towards leopards. Although available leopard habitat has likely increased in parallel with the growth of the commercial wildlife industry (Thorn, et al., 2011), it is unlikely that trophy hunting of leopards directly incentivizes the private sector to conserve leopard habitat.

It is hoped that the adaptive management framework recently adopted in South Africa, through which trophy hunting is limited to areas with stable (or increasing) leopard populations as demonstrated by scientifically robust data on leopard population trends, will encourage collaborative landowner participation in the South African Leopard Monitoring Project, and ultimately incentivize management practices that contribute towards the conservation of leopards.

8. Livelihood/ socio-economic benefits and impacts

In South Africa the promulgation of the Game Theft Act (No. 105 of 1991, as amended in Acts 18 of 1996 and 62 of 2000), which grants conditional ownership of wildlife to private landowners who obtain a Certificate of Adequate Enclosure (CAE), consolidated the foundations of an economically viable wildlife industry (Carruthers, 2008). The right of ownership of wildlife, combined with a growing understanding that wildlife ranching was ecologically and financially sustainable, along with significantly reduced subsidies for conventional agriculture and increasing financial incentives for commercial wildlife ranching, has led to a tremendous increase in land under wildlife and game numbers over the past 30 years, and the establishment of a formal wildlife sector in South Africa.

Southern Africa has a particularly well-established sport hunting industry that generates substantial revenues. Lindsey, et al. (2007) estimated the annual revenues generated by trophy hunting in sub-Saharan Africa to be approximately USD200million, with USD100 million of those accrued by South Africa. Trophy hunters in South Africa were reported to spend an average of USD20,136 per hunter for the 2015/2016 hunting season, in total USD131 million for the approximately 7600 international hunters hosted by South Africa in that year (TREES, 2017). In addition to direct financial values, the Professional Hunters’ Association of South Africa (PHASA) estimates that thousands of jobs are created by professional hunting in South Africa, including hunting outfitters, professional hunters and other jobs created by international hunting tourists. Moreover, >60 000 bed nights were booked by international hunting tourists in 2014, amounting to approximately USD9 million (Sinovas, et al., 2016).



The total estimated financial value of leopard trophy exports from SADC (the Southern African Development Community) over the period 2005 – 2014 was USD9 964 702 (USD1520/leopard trophy) (Sinovas et al., 2016). Approximately 14% of these trophies originated in South Africa (USD1 395 058). Leopard is also the most valuable trophy exported from SADC, followed by elephant (USD1303) and hippopotamus (USD759) (Sinovas, et al., 2016).

9. Research projects underway

Research from South Africa, for the period 1982 – 2012, accounted for a significant proportion of the peer-reviewed literature on leopards; more than double that of any other range state. However, unlike leopard research conducted elsewhere, South African studies focused primarily on ecological themes, rather than conservation and management (Balme,et al., 2013). Current research projects within South Africa include:

  • Panthera: Furs for Life – combatting the illegal trade in leopard skins for cultural regalia through education, policy and the provision of faux leopard furs.
  • Panthera: The origins of leopard skins entering Shembe markets.
  • Landmark Leopard and Predator Foundation: ecology of leopards, remedial action forinjured leopards, and conflict management with livestock owners.
  • Primate and Predator Project: conducting research into the status of leopards outside of protected areas and in the Soutpansberg Mountains, Limpopo Province.
  • North West Leopard Project: investigating the ecology of leopards in the province through camera trapping and GPS collars, with a view to enable province-wide management (e.g. setting quotas, conflict management and translocation appraisal).
  • Cape Leopard Trust: continuing work on leopards in the greater Western Cape, and to venture into Northern Cape. Farmer education and ecological research.
  • Mpumalanga Tourism and Parks Agency: Ingwe Leopard Project: Greater Lydenburg area; Kruger National Park western boundary carnivore monitoring, including the neighbouring rural areas; spatial ecology, habitat utilisation, population demographics and conservation of leopards in the Loskop Dam Nature Reserve10. Conclusion and recommendationsLeopards are generally considered uncommon in South Africa, however estimates of the size of the national population vary so widely as to make them meaningless. Estimated leopard densities in South Africa range from 0.2 ± 0.0 leopards/100 km2 to 12.2 ± 2.4 leopards/100 km2. South Africa’s leopard population is apparently declining by 8% per year and significant declines in leopard density have been observed in some areas. Some stable populations appear to be well below their potential capacities, while other areas with prime leopard habitat seem to no longer have functioning leopard populations. At present, the illegal killing of leopards for skins and other body parts for traditional ceremonies and



medicines is believed to be the major threat facing leopard within South Africa. Addressing the illegal skin trade remains a compliance and enforcement challenge. In some provinces, progress has been made in reducing the number of leopards killed as DCAs, though the recent shift in South Africa’s game ranching industry to high-value species and colour variants has likely decreased tolerance towards leopards and increased levels of retaliatory killing.

Both national and provincial legislation provides for the protection and conservation of leopard in South Africa. Since the publication of a non-detriment finding (NDF) on leopard in December 2015, the government of South Africa has made concerted efforts to implement measures recommended in the NDF in order to improve the management of the species:

  1. a)  The South African Leopard Monitoring Project has been established, providing for a standardised, rigorous framework using systematic camera-trap surveys for the monitoring of leopard population trends throughout South Africa.
  2. b)  An adaptive approach for the management of trophy hunting has been adopted, and population trend data generated by the South African Leopard Monitoring Project are being used to inform leopard hunting quotas on an annual basis. Hunting will be restricted to leopard hunting zones where scientifically robust data on leopard density trends indicate overall stable (or increasing) populations.
  3. c)  Norms and standards for the trophy hunting of leopard in South Africa are currently under development, the general approach of which is to: 1) distribute hunting effort evenly across leopard range in South Africa; 2) limit hunting to male leopards of at least seven years of age; and 3) ensure the mandatory submission of hunt return data and trophy photographs following all leopard hunts (both successful and unsuccessful).
  4. d)  An online application and submission system for the management of trophy hunting has been developed and roll out to the provinces is imminent. Hunt return data will be used to determine catch-per-unit-effort and harvest composition as indices of relative abundance, while the data capture for leopards killed as DCAs will improve monitoring of DCA off-take.

Between 2005 and 2016, South Africa did not fully utilize its export quota of 150 leopard hunting trophies and skins for personal use, having exported on average 73 per year, mostly as trophies. It is therefore unnecessary to consider an increase in the export quota. On the other hand, a reduction in the export quota would limit the flexibility that is crucial for the adaptive management approach adopted by South Africa for the allocation of leopard hunting quotas. Considering that the leopard is the most valuable hunting trophy exported from SADC, it is hoped that this adaptive management approach will encourage collaborative landowner participation in the South African Leopard Monitoring Project, and ultimately incentivize management practices that contribute towards leopard conservation.

It is therefore recommended that South Africa’s export quota for leopard hunting trophies and skins for personal use (Resolution Conf. 10.14 (Rev. CoP16)), as originally informed by a Population and Habitat Viability Analysis (PHVA), be retained at 150.




Balme G., Dickerson, T., Fattebert, J., Lindsey, P., Swanepoel, L., and Hunter, L., 2014. Ethics and wildlife studies: the questionable use of radio-telemetry in leopard research.Conservation Letters 7(1); 3-11.

Balme, G.A., Lindsey, P.A., Swanepoel, L.H., and Hunter, L.T.B., 2014. Failure of Research to Address the Range wide Conservation Needs of Large Carnivores: Leopards in South Africa as a Case Study. Conservation Letters 7, 3-11.

Balme, G.A., Batchelor, A., de Woronin Britz, N., Seymour, G., Grover, M., Hes, L., Macdonald, D.W., and Hunter, L.T.B., 2013. Reproductive success of female leopardsPanthera pardus: the importance of top-down processes. Mammal Review 43, 221-237.

Balme, G.A. and Hunter, L.T., 2013. Why leopards commit infanticide.Animal

Behaviour86(4), pp.791-799.

Balme G. A., Hunter L., and Braczkowski A. R., 2012. Applicability of Age-Based Hunting

Regulations for African Leopards. PLoS ONE 7(4): e35209. doi:


Balme, G., Slotow, R. and Hunter, L., 2010a. Edge effects and the impact of non‐protected areas in carnivore conservation: Leopards in the Phinda–Mkhuze Complex, South Africa.Animal Conservation 13(3); 315-323.

Balme, G., Hunter, L., Goodman, P., Ferguson, H., Craigie, J. and Slotow, R., 2010b. An adaptive management approach to the trophy hunting of Leopards: a case‐study from Kwa‐Zulu Natal, South Africa. Pages 341‐352 in ‘The Biology and Conservation of Wild Felids’, D. Macdonald & A. Loveridge (eds.), Oxford University Press, New York.

Balme, G., Slotow, R. and Hunter, L., 2009. Impact of conservation interventions on the dynamics and persistence of a persecuted Leopard (Panthera pardus) population.Biological Conservation, 142: 2681‐2690.

Carruthers, J. 2008. “Wilding the farm or farming the wild”? The evolution of scientific game ranching in South Africa from the 1960s to the present. Transactions of the Royal Society of South Africa 63:160-181.

Chase Grey, J.N., Kent, V.T., & Hill, R.A., 2013. Evidence of a High Density Population of Harvested Leopards in a Montane Environment. PLoS ONE 8(12): e82832. doi:10.1371/journal.pone.0082832

Chase Grey, J.N., Bell, S., and Hill, R.A., 2017. Leopard diets and landowner perceptions of human wildlife conflict in the Soutpansberg Mountains, South Africa. Journal for Nature Conservation 37, 56–65.

CITES Trade Database, UNEP World Conservation Monitoring Centre, Cambridge, UK. Constant, N.L., 2014. A socio-ecological approach towards understanding conflict between leopards (Panthera pardus) and humans in South Africa: Implications for leopard conservation and farming livelihoods. PhD Thesis, Department of Anthropology, Durham

University, USA.
Crookes, K.R., Sanjayan, M.A., & Doak, D.F. 1998. New insights on cheetah conservation

through demographic modelling. Conservation Biology 12: 889-895.
Dickson, B., Hutton, J. and Adams, B., 2009. Recreational Hunting, Conservation and Rural

Livelihoods: Science and Practice Wiley-Blackwell, Oxford, UK. 384 pp. 14

Chapman, S. & Balme, G., 2010. An estimate of leopard population density in a private

reserve in KwaZulu-Natal, South Africa, using camera-traps and capture-recapture

models. South African Journal of Wildlife Research 40.2 (2010): 114-120.


Di Minin, E., Hunter, L.T.B., Balme, G.A., Smith, R.J., Goodman, P.S., & Slotow, R., 2013. Creating larger protected areas enhancing the persistence of big game species in the Maputaland-Pondoland-Albany biodiversity hotspot. PLoS ONE 8, e71788.

Friedmann, Y. & Traylor‐Holzer, K., 2008. Leopard (Panthera pardus) case study. NDF Workshop Case Studies, Mexico 2008.

Gaona, P., Ferreras, P., & Delibes, M., 1998. Dynamics and viability of a metapopulation of the endangered Iberian lynx. Ecol Mono 68: 349-370.

Hunter, L., Henschel, P. & Ray, J., 2013. Panthera pardus. In: The Mammals of Africa. Vol. V: Carnivores, Pangolins, Equids and Rhinoceroses (Ed. by J. Kingdon & M. Hoffmann), pp. 159-168. London: Bloomsbury

IUCN. 2016. Informing decisions on trophy hunting: a briefing paper for European Union decision-makers regarding potential plans for restriction of imports of hunting trophies. IUCN Briefing Paper April 2016.

Jacobson, A.P., Gerngross, P., Lemeris Jr, J.R., Schoonover, R.F., Anco, C., Breitenmoser- Würsten, C., Durant, S.M., Farhadinia, M.S., Henschel, P., Kamler, J.F. & Laguardia, A., 2016. Leopard (Panthera pardus) status, distribution, and the research efforts across its range. PeerJ 4, e1974.

Jones, J.L. 2006. Transboundary conservation in Southern Africa: Exploring conflict between local resource access and conservation, PhD Thesis. University of Pretoria, Pretoria.

Lindsey, P., 2011. A review of the consumptive utilization and trade of Cheetahs and Leopards in South Africa. Endangered Wildlife Trust report.

Lindsey, P. A., P. A. Roulet, and S. S. Romanach. 2007. Economic and conservation significance of the trophy hunting industry in sub-Saharan Africa. Biological Conservation134:455-469.

Lindsey, P., du Toit, J.T. & Mills, M.G.L., 2005. Attitudes of ranchers towards African wild dogs Lycaon pictus: conservation implications for wild dogs on private land. Biological Conservation 125: 113‐121.

Martin, R. B. and De Meulenaer, T., 1988. Survey of the status of the Leopard (Panthera pardus) in subSaharan Africa. Convention on the International Trade of Endangered Species Secretariat, Lausanne, Switzerland.

Martins, Q. and Martins, N., 2006. Leopards of the Cape: conservation and conservation concerns. International Journal of Environmental Studies, 63(5), 579–585.

Mann, G., Pitman, R., Whittington-Jones, G., Thomas, L., Broadfield, J., Taylor, J., Rogan, M., and Balme, G.,

2017. South African Leopard Monitoring Project. Annual report on

monitoring activities in 2017. Report to the Scientific Authority of South Africa, November


Maputla, N. W., Chimimba, C. T., and Ferreira, S. M., 2013. Calibrating a camera trap–

based biased mark–recapture sampling design to survey the leopard population in the

N’wanetsi concession, Kruger National Park, South Africa.African Journal of

Ecology51(3), 422-430.

McManus, J.S., Dalton, D.L., Kotze, A., Smuts, B., Dickman, A., Marshal, J.P., and Keith, M., 2015. Gene flow and population structure of a solitary top carnivore in a human- dominated landscape. Ecology and Evolution 5(2): 335-344.

Packer, C., Brink, H., Kissui, Maliti, H., Kushnir, H. and Caro, T., 2011. Effects of trophy hunting on lion and leopard populations in Tanzania. Conservation Biology, 25: 142‐153.

Peebles, K.A., Wielgus, R.B., Maletzke, B.T., and Swanson, M.E., 2013. Effects of Remedial Sport Hunting on Cougar Complaints and Livestock Depredations. PLoS ONE 8, e79713.



Pirie, T.J., Thomas, R.L., & Fellowes, M.D.E., 2017. Increasing game prices may alter farmers’ behaviours towards leopards (Panthera pardus) and other carnivores in South Africa. PeerJ 5:e3369; DOI 10.7717/peerj.3369

Pitman, R.T., Fattebert, J., Williams, S.T., Williams, K.S., Hill, R.A., Hunter, L.T.B., Slotow, R., and Blame, G.A., 2016. The conservation costs of game ranching. Conservation Letters; doi:10.1111/conl.12276

Pitman, R.T., Swanepoel, L.H., Hunter, L., Slotow, R., and Balme G.A., 2015. The importance of refugia, ecological traps and scale for leopard population management.Biodiversity and Conservation 24(8), 1975-1987.

Sinovas, P., Price, B., King, E., Davis, F., Hinsley, A., Pavitt, A., and Pfab, M. 2016.Southern Africa’s wildlife trade: an analysis of CITES trade in SADC countries. Technical report prepared for the South African National Biodiversity Institute (SANBI). UNEP- WCMC, Cambridge, UK.

Swanepoel, L.H., Somers, M.J., van Hoven, W., Schiessmeier, M., Owen, C., Snyman, A., Martins, Q., Senekal, C., Camacho, G., Boshoff, W., and Dalerum, F., 2014. Survival rates and causes of mortality of leopards Panthera pardus in southern Africa. Oryx doi: 10.1017/S0030605313001282

Swanepoel, L.H., Lindsey, P., Somers, M.J., van Hoven, W., and Dalerum, F., 2014. The relative importance of trophy harvesting and retaliatory killing of large carnivores: South African leopards as a case study. South African Journal of Wildlife Research 44(2): 115- 134.

Thorn, M., Green, M., Scott, D., and Marnewick, K., 2013. Characteristics and determinants of human-carnivore conflict in South African farmland. Biodiversity Conservation 22: 1715-1730.

TREES. 2017. A marketing and spending analysis of trophy hunters 2015/2016 season., Tourism Research in Economic Environs and Society, North-West University, Potchefstroom, South Africa.

UNEP-WCMC, 2013. Assessing potential impacts of trade in trophies imported for hunting purposes to EU-27 on conservation status of Annex B species. Part 2: Discussion and case studies. Prepared for the European Commission by UNEP-WCMC. Cambridge, UK. 34 pp.

R. Soc. open sci. 4: 161090. http://dx.doi.org/10.1098/rsos.161090.16

Ramesh, T., Kalle, R., Rosenlund, H., and Downs, C.T., 2017. Low leopard populations in

protected areas of Maputaland: a consequence of poaching, habitat condition, abundance

of prey and a top predator. Ecology and Evolution 7(6); 1964-1973

St John, F. A., Keane, A. M., Edwards-Jones, G., Jones, L., Yarnell, R. W., and Jones, J. P.,

2011. Identifying indicators of illegal behaviour: carnivore killing in human-managed

landscapes. Proceedings of the Royal Society B: Biological Sciences, rspb20111228.

Swanepoel, L. H., Lindsey, P., Somers, M. J., Hoven, W. V., and Dalerum, F., 2013. Extent

and fragmentation of suitable leopard habitat in South Africa. Animal Conservation16(1),


Thorn, M., Green, M., Keith, M., Marnewick, K., Bateman, P. W., Cameron, E. Z., and Scott,

D. M., 2011. Large-scale distribution patterns of carnivores in northern South Africa:

implications for conservation and monitoring. Oryx, 45(04), 579-586.

Treves, A., and Bruskotter, J. (2014). Tolerance for Predatory Wildlife. Science (New York,

NY), 344(6183), 476-477.

Williams, S.T., Williams, K.S., Lewis, B.P., and Taylor R. 2017. Population dynamics and

threats to an apex predator outside protected areas: implications for carnivore