Red List of South African Species

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Vulnerable (VU)
Assessors: Lourens Swanepoel Williams Samuel John Power Andrei Snyman Ian Gaigher Charl Senekal Quinton Martins
Contributors: Samantha Page Michael Hoffmann Gerrie Camacho
Reviewers: Matthew Child Guy Balme

Rationale

Although this is a widespread species within the assessment region, it is secretive and faces severe threats, especially outside protected areas. The most systematic population estimate ranges from 2,813–11,632 Leopards, which equates to 1,688–6,979 mature individuals (60% mature population structure). All subpopulations number fewer than 1,000 mature individuals except the bushveld subpopulation (Kruger National Park, Limpopo, Mpumalanga and North West Province), which is likely to number between 1,113–4,454 mature individuals. However, as these estimates are derived from habitat suitability models and could under- or over-estimate actual abundances, caution should be used when applying the numbers (for example, in setting hunting quotas). Two independent simulation models project an ongoing population decline, largely due to unsustainable rates of persecution (direct and indirect) and a poorly managed trophy hunting industry, over the next 25 years. This is corroborated by empirical research that shows Leopard proportional survival in non-protected areas is only 57 ± 14% compared to 86 ± 5% inside protected areas. Additionally, Leopards within the assessment region are facing an emerging threat of being hunted for cultural regalia (for example, an estimated 17,240–18,760 illegal Leopard skins are believed to be used for ceremonial church activities). Similarly, the rise of intensive wildlife breeding for high-value game species may also be increasing the extent and intensity of persecution.

Province-wide population ecology studies at four independent localities (two in KwaZulu-Natal Province and two in Limpopo Province) show severe declines over short time periods: Leopard density has declined far in excess of 10% over the past 4–7 years (Welgevonden: 33% decline in 6 years; Lajuma: 22% decline in 7 years; Hluhluwe-iMfolozi: 40% decline in 4 years; uMkhuze: 13% decline in 5 years; G. Balme unpubl. data), which suggests that the data are not local aberrations or fluctuations but indicate systemic or widespread population decline.

Thus we list Leopards as Vulnerable C1 due to small population size and an estimated continuing decline of at least 10% over three generations (18–27 years). Severe declines are corroborated both by model simulations and empirical data from disparate geographical locations within protected areas and private land. Given the lower survival rate of Leopards outside protected areas, declines are suspected to be similar or more severe in such areas. Although the rate of decline between this assessment and the previous assessment is difficult to measure due to the variance in population estimates, we construe the net continuing population decline, the emerging threat of being hunted for cultural regalia, and the possible increase in persecution with wildlife ranch expansion as a genuine change in listing. Monitoring frameworks, which enable provinces to track regional Leopard subpopulation trends should be established so as to more accurately estimate population reduction over three generations, as this species may justify a more threatened listing under the A criterion. Key interventions include the adoption of sound harvest management regulations, the use of livestock guarding dogs to reduce conflicts, and the use of faux Leopard skins as a cultural regalia substitute.

Regional population effects: Although no quantitative assessment has been done regarding the extent of suitable Leopard habitat across the southern African region, dispersal occurs between neighbouring countries (Fattebert et al. 2013). This has to some extent been facilitated by the establishment of Transfrontier Parks. However, continued human population growth and livestock/game farming along South African and neighbouring country borders (even within some Transfrontier Parks), means the associated Leopard-landowner conflict might limit the rescue effect of South Africa's neighbouring countries (Purchase & Mateke 2008). Furthermore, Leopard subpopulations along South African borders also face similar threats like illegal harvesting, persecution, poorly managed trophy hunting and incidental snaring (Purchase & Mateke 2008; Jorge et al. 2013). As such, although the rescue effect is possible, it is unlikely to be a significant factor in reducing extinction risk within the assessment region.

Distribution

Leopards remain widely, but patchily, distributed (Stein et al. 2016), having been lost from at least 37% of their historical range in sub-Saharan Africa (Ray et al. 2005), and 28–51% of their historical range in Southern Africa (Jacobson et al. 2016). The most marked range loss has been in the Sahel belt, as well as in Nigeria, Malawi and, importantly, South Africa (Stein et al. 2016), where Leopards have become extinct in 67% of the country (Jacobson et al. 2016). The species has become locally extinct in areas of high human density or extensive habitat transformation (Hunter et al. 2013). Within the assessment region, they range extensively across all provinces (except the Free State Province and the greater Karoo basin in the Northern and Western Cape provinces), including Swaziland but not Lesotho; and they occur in all biomes of South Africa, with a marginal occurrence in the Nama Karoo and Succulent Karoo biomes. While Leopards were present in both Free State Province and Lesotho historically (Lynch 1983, 1994), they are very rare or absent entirely from these areas today. However, there has been a recent record (2014) from Clocolan, Free State Province by the Department of Economic Development, Environment, Conservation and Tourism, that occasionally attends to Leopards as damage-causing animals (N. Collins pers. comm. 2016). Additionally, Swanepoel et al. (2014) estimated that 8–26 individuals may occur in the Free State Province based on habitat suitability.

Available habitat is becoming increasingly rare: recent habitat suitability models classed only around 20% of South Africa as suitable habitat (Swanepoel et al. 2013), although both the extent of occurrence and area of occupancy inferred to have remained stable or even increased from 2000 to 2010 in North West Province (Thorn et al. 2011; Power 2014). All areas identified as suitable habitat need to be surveyed for confirmed Leopard presence. For example, potential Leopard habitat in the Western Cape, excluding isolated areas where they have been sighted, is approximately 40,000 km2 (Martins 2010), composed of 10,000 km2 conserved areas, state land and mountain catchment areas (prime habitat), with the remaining 30,000 km2 comprising crop and livestock farming and small towns, which may support resident Leopards or transitory individuals. Further research and field surveys investigating spatial patterns of Leopard subpopulations outside of protected areas is needed (Balme et al. 2014). They are able to disperse large distances. For example, an individual from Maputaland, KwaZulu-Natal Province, traversed three countries covering 353 km (Fattebert et al. 2013).

Suitable Leopard habitat in South Africa has been further fragmented into four core areas, based on MaxEnt models using true positive data (Swanepoel et al. 2013), namely 1) the west coast and southeast coast of the Western and Eastern Cape Provinces; 2) the interior of KwaZulu-Natal Province; 3) the Kruger National Park and the interior of Limpopo, Mpumalanga and North West Provinces; and 4) the northern region, containing the Kgalagadi Transfrontier Park (KTP) and adjacent areas of the Northern Cape and North West Provinces. There may be a fifth subpopulation pending further investigation: we are still unsure of the Northern Cape population documented in Namaqualand and up to the Richtersveld, which may be connected to the Western Cape subpopulation, but phenotypically they are more similar to Leopards elsewhere in the country (McManus et al. 2015a).

Population trend

Trend

The Leopard is an adaptable, widespread species that nonetheless may have many threatened subpopulations. Leopard population size and trends are notoriously difficult to estimate, due to their secretive nature and the high financial costs involved in population monitoring. As such, density and population estimates can have low precision which makes interpretation difficult. The most systematic estimate (based on habitat availability and suitability, as well as a range of density estimates for each province) ranges from 2,813–11,632 Leopards, with a median (best scenario) estimate of 4,476 (Swanepoel et al. 2014). If we assume that 60% of the population is mature, (30% males, 30% females, 15% sub-adult males and females and 10% juveniles; Swanepoel et al. 2014), there are 1,688–6,979 mature individuals within the assessment region. This estimate is similar to the 4,250 estimated by Daly et al. (2005), but much lower than the 10,000 estimated by Martin and De Meulenaer (1988). The latter has been criticised as being an overestimate (Norton 1990). Such large variance makes quantitative interpretation difficult and thus these data can only be used as a rough guideline of the South African Leopard population. Caution should therefore be applied when using these data quantitatively (for example, to set hunting quotas). The estimated generation length ranges from six years (IUCN unpubl. data) to nine years (Pacifici et al. 2013), yielding the three generation window as 18–27 years. Leopards become sexually mature at 2.5–3 years old (Nowell & Jackson 1996; Skinner & Chimimba 2005). First-year mortality was estimated to be 41–50% (Martin & De Meulenaer 1988; Bailey 1993) and cub survival was estimated to be only 37% (Balme et al. 2013).

Two independent models indicate a continuing decline in the population: Daly et al. (2005) projected a continuing population decline of 16% between 2005 and 2025 under a trophy hunting quota of 150 animals per year. This is congruent with stochastic population results from Swanepoel et al. (2014), where all provincial populations showed consistent declines under a range of realistic scenarios of harvest and damage-causing animal control over the next 25 years, although overall extinction risk is low (< 10% probability). Furthermore, current province-wide population ecology studies in KwaZulu-Natal and Limpopo Provinces show subpopulation declines in several protected areas (Welgevonden: 33% decline in 6 years; Lajuma: 22% decline in 7 years; Hluhluwe-iMfolozi: 40% decline in 4 years; uMkhuze: 13% decline in 5 years; G. Balme et al. unpubl. data). Additional survey data reveal negative subpopulation trends (from 2–4 repeated surveys) for 10 out of 13 sampled protected areas across the country (G. Balme unpubl. data). Thus, declines projected from model outputs are corroborated by empirical data from localities in different regions, which 1) suggests a generalised widespread decline (rather than local short-term fluctuations or aberrations); and 2) a rate of decline that exceeds the 10% threshold over three generations and may indicate a greater national population reduction. Further monitoring is required to estimate or project past and future population reduction.

We suspect overall population declines may be greater, as estimated declines are from within protected areas and Leopard survival is lower outside of protected areas, and hence may be subject to more severe declines. For example on private land in the western Soutpansberg Mountains, Limpopo Province, preliminary analyses suggest that the Leopard density has declined from 10.7 individuals / 100 km² in 2008 (Chase Grey et al. 2013) to approximately 4 Leopards / 100 km² in 2015 (S. Williams unpubl. data). Research shows Leopard survival in non-protected areas is 57 ± 14% while in protected areas is 86 ± 5% (Swanepoel et al. 2015b). Survival is especially low for females (Balme et al. 2013; Swanepoel et al. 2014). Similarly, densities can be low in highly suitable areas, whether protected or not. For example, in the Phinda–Mkuze complex (KwaZulu-Natal Province) Leopard density declined from the core of the reserve (11 ± 1 individuals / 100 km2) to the border (7 ± 1 individuals / 100 km2), being the lowest in non-protected areas adjoining the reserve (3 ± 0.9 individuals / 100 km2), and was not related to prey abundance or interspecific competition (Balme et al. 2010b). In North West Province, there is an estimated annual persecution rate of 0.4 individuals killed / 100 km2 (Thorn et al. 2012), while the western Kalahari region may have experienced a significant decline in Leopard numbers as inferred from the number of damage-causing animal control records (Power 2014).

Recent multiscale genetic analysis suggested southern African Leopards comprise a single population of distinct geographically isolated groups (Ropiquet et al. 2015). This supports previous analyses (Miththapala et al. 1996; Uphyrkina et al. 2001), and confirms evidence for geographically isolated groups (for example, Western and Eastern Cape are geographically isolated from Limpopo; Tensen et al. 2011). However, genetic work from Leopard subpopulations within Eastern and Western Cape provinces detected significant population clustering, with low emigration and immigration between subpopulations (McManus et al. 2015a, Swanepoel et al 2013). Evidence is thus amassing that suggests local population isolation can be attained within relatively few generations highlighting the importance of management actions that aims to increase habitat connectivity and reduce human–carnivore conflict (McManus et al. 2015a).

Threats

Within the assessment region, the major threats to Leopards are intense persecution, both directly through hunting (trophy or DCA control) or indirectly through snaring, and demand for their skins for cultural regalia (Hunter et al. 2013). Compared to other African countries, South Africa is highly developed and thus Leopard subpopulations have become fragmented (Swanepoel et al. 2013), and there has been a long history of persecution owing to real or perceived livestock depredations (Stuart 1981; Norton 1986; Lindsey et al. 2005; St John et al. 2011). These threats are more pronounced outside protected areas (Swanepoel et al. 2015a,b), where mortality on non-protected land is due to legal and illegal damage hunting/control, whereas snaring and poisoning are significant causes of mortality inside protected areas. For example, in the Soutpansberg Mountains, Limpopo Province, the most common cause of death of eight Leopards that were fitted with GPS collars between 2012 and 2015 was snaring, followed by illegal activity to protect livestock predation, such as shooting and poisoning (S. Williams unpubl. data). North West authorities, however, do mitigate this by removing snares from collared Leopards by airborne immobilisation, treatment and re-release. Stationary Leopards are thus immediately investigated with the suspicion that they are ensnared (R.J. Power pers. obs. 2015).

Ongoing habitat loss and fragmentation also threatens the recovery of this species (see below).
  1. Direct persecution: Sustainability of trophy harvest is reduced due to high incidences of direct persecution in South Africa (Swanepoel et al. 2014). This, on its own, reduces Leopard population size and disrupts social organisation (Balme et al. 2013). Swanepoel et al. (2014) estimated that 35% of all Leopards killed in retaliatory actions are reproductive females. Such removals of females leads to reduced survival of Leopards in non-protected areas and thus affects long term population viability (Swanepoel et al. 2015b). Compounding this problem is an obvious lack of clear national conservation objectives resulting in large disparity in the number of DCA permits issued in different provinces, which ranges from 17 in one year to only two between 2007 and 2013 in the same province (Q. Martins and R.J. Power pers. obs. 2015). The reduction in permits was partly attributed to a local conservation NGO monitoring the permit issuing process. Direct poaching of Leopards is also suspected to be increasing due to the demand for skins (see below), which may be far more severe a threat than problem-animal control and unsustainable trophy hunting combined.
  2. Cultural regalia: There appears to be a strong demand for Leopard skins for cultural regalia. Preliminary capture-recapture analyses suggest that members of the Nazareth Baptist Church (also known as the Shembe) may be in possession of 17,240–18,760 illegal Leopard skins with a subsequent high rate of removal from the wild (G. Balme unpubl. data). This represents an emerging threat to this species within the assessment region.
  3. Trophy hunting: Unsustainable and poorly-managed trophy hunting can cause subpopulation decline (Balme et al. 2009, 2010b; Packer et al. 2011; Swanepoel 2013; Swanepoel et al. 2014). Poorly managed trophy quotas are characterized by the hunting of females, clumped harvests (excessive hunting around protected areas, and multiple tags in the same area) and hunting of inappropriate age classes (for example, excessive hunting of prime adults males) (Packer et al. 2009, 2011; Balme et al. 2012). Excessive and clumped harvest of male Leopards < 7 years old can destabilise Leopard social organisation, leading to reduced cub survival and increased female mortality (Balme et al. 2012). Similarly, hunting females can reduce overall reproductive output causing population declines (Dalerum et al. 2008). While hunting of females is detrimental, South African law still allows for female harvest (32–50% of hunted Leopards are females; Swanepoel et al. 2014). Even in countries with only male harvest, like Tanzania, females comprised 29% of 77 trophies shot between 1995 and 1998 (Spong et al. 2000). The use of national population estimates to set trophy hunting quotas is perilous. For example, the over-estimate of 10,000 Leopards in South Africa (Martin & De Meulenaer 1988; Norton 1990) was used by conservation authorities to set hunting quotas from 2005 onwards (Daly et al. 2005).
  4. Indirect persecution: While snares laid out for bushmeat hunting threaten Leopards, especially inside protected areas, a rapidly increasing threat is the poisoning of carcasses, either as a means of predator control or incidentally. The rise of intensive wildlife breeding for high-value game species may also be increasing the extent of both direct and indirect persecution (Thorn et al. 2012, 2013).
  5. Radio-collars: Another significant and localised threat is the injudicious use of radio-collars for research and recreational purposes. Sub-adults exhibit rapid growth and have a high neck-head circumference ratio (G. Balme unpubl. data). Collars can asphyxiate Leopards if they cannot be loosened. Poor capture techniques also pose a threat to Leopards. Despite this, radio-collars are widely deployed on Leopards in South Africa, often with little oversight. Eighty percent of Leopard projects in South Africa, reviewed in a recent study, used radio-telemetry (N = 39; G. Balme unpubl. data.), and most of these projects failed to deploy breakaway devices on collars. Similarly, many (63%) did not contribute to the scientific literature, even though some were active for over 12 years (Balme et al. 2014). It appears the motivation for much Leopard research in South Africa, particularly hands-on research such as radio-collaring, is to enable commercial volunteer programmes, where laypeople (typically foreign graduate students) pay to experience research (Balme et al. 2014). North West Province have instituted the policy of controlling all Leopard collaring under their bannership and research, and insist upon recaptures and collar removal via conditioning animals upon recapture.
  6. Road collisions: Although the effect of this threat on the population is unknown, Leopards are amongst the species killed on roads, even within protected areas.

Uses and trade

Leopards are hunted (legally and illegally) as a trophy animal within the assessment region. When properly managed, trophy hunting should have little effect on population persistence (Swanepoel et al. 2014); however, research from KwaZulu-Natal (Balme et al. 2009, 2010b), Limpopo (Pitman et al. 2015) and Tanzania (Packer et al. 2011) suggest that poorly managed trophy hunting might lead to Leopard population declines . In South Africa, population models, which only include off-take from trophy hunting, suggest that current trophy harvest levels have little impact on population persistence (Swanepoel 2013; Swanepoel et al. 2014). However, when other forms of human-induced mortality (for example, legal and illegal retaliatory killing due to human-Leopard conflict) are included, trophy hunting quotas become unsustainable (Swanepoel 2013; Swanepoel et al. 2014). The detrimental impacts of trophy hunting may be reduced by improving current management practices, most notably by banning the hunting of female Leopards and ensuring the equitable distribution of hunting effort across Leopard range (Balme et al. 2010a). Suspicions are that captive-bred-animals are laundered into the trophy hunting industry and this should also be investigated. Additionally, there is a suspected industry in catching Leopards from the wild and providing them for the trophy hunting industry. Authorities are constantly confiscating such animals and attempting to repatriate them elsewhere without any knowledge of their origin (R.J. Power unpubl. data).

The likely impacts of the illegal skin trade also need to be factored into assessments of harvest sustainability. Surveys suggest as many as 17,240–18,760 illegal Leopard skins are used by members of the Shembe Church for religious regalia and may be replaced every 3–5 years due to wear (G. Balme unpubl. data). Although interviews with traders suggest many skins originate outside South Africa (G. Balme unpubl. data), the trade is likely to affect Leopard population viability throughout the assessment region. While wildlife ranching and game farming might be increasing in suitable habitat for Leopards (Power 2014), such industries are normally in conflict with predators (Thorn et al. 2012), especially as the recent shift to breeding high-value species and colour variants has increased hostility towards carnivores (Thorn et al. 2013). This is reflected by the rapid increase in the number of damage-causing animal (DCA) permit applications received from game farms in areas such as the Waterberg (Lindsey et al. 2011). An increase in ranching rare/expensive game, especially intensive breeding for trophies and colour variants, may thus impact negatively on the Leopard population through increased persecution, exclusion with predator-proof fences and limitation of gene flow. Such conflict results in two outcomes; 1) increased persecution or legal removal (DCA permits) and 2) possible exclusion through improved fencing to keep predators out. While Leopards appear to be unhindered by standard game farm fences (Fattebert et al. 2013), the quality of predator-proof fencing has improved to such an extent that it may hinder their movement between properties (du Plessis & Smit 1999).

Conservation

Although Leopards occur in numerous protected areas across their range, the majority of the population occurs outside of protected areas, necessitating a need for improved conflict mitigation measures, trophy hunting management, non-lethal mitigation actions, centralised monitoring of trophy harvest and quality, issuing of DCA permits as well as providing education programmes to ensure Leopards do not become locally threatened. Currently, the best conservation effect on Leopard conservation in South Africa can be made along two general fronts, namely policy development and conflict mitigation. These can be distilled into four pillars of conservation action: 1) livestock and game conflict mitigation, 2) applying sustainable trophy hunting regulations, 3) reducing the illegal trade in skins and 4) protected area expansion to create a more resilient population overall.

Firstly, it is important to revise current Leopard management policies implemented by local conservation authorities and councils. These include adoption of stringent control methods in Leopard trophy hunting, which include age based harvest (for example, harvest of old males; Balme et al. 2010a, 2012), enforcement of male only harvest (Balme et al. 2010a; Swanepoel et al. 2014) and non-spatial clumped harvest (Balme et al. 2010a). Furthermore, issuing permits to destroy damage-causing Leopards needs to be revised by including better conflict mitigation actions (for example, guard dogs and predator-proof fencing; McManus et al. 2015b), and implementing/monitoring mitigation actions, especially preventing the destruction of female Leopards (Swanepoel et al. 2014). Secondly, such policies being implemented by the different provincial governments must be continuously monitored. These two management actions should be nationally implemented. A national monitoring framework should be established to analyse trends in Leopard off-take (via the different mortality sources) and to detect changes in Leopard occupancy and local densities. However, regional populations have additional threats which should be addressed at a local scale. Thus, conservationists should focus on the following interventions:
  • Livestock conflict mitigation through use of guarding dogs and improved livestock husbandry (Marker et al. 2005). While various pilot projects have been established in Limpopo, North West, Northern and Western Cape provinces, little research has been done about their overall effectiveness within the assessment region, especially for Leopards. Preliminary findings suggest that livestock guarding dogs can decrease depredation by 69% (McManus et al. 2015b).
  • Applying and enforcing more sustainable trophy hunting and damage-causing animal regulations (Balme et al. 2012), which are described above.
  • Reducing the illegal trade in skins by providing faux furs for use at cultural ceremonies. Since the project began in 2013, 5,160 Leopard skins have been donated by the conservation NGO Panthera to members of the Shembe Church. Results are preliminary, but the ratio of fake to authentic skins observed at Shembe gatherings has increased from 1:8 to 1:4 (G. Balme et al. unpubl. data). However, the overall impact of the intervention on the regional Leopard population remains unknown. More generally, this also includes interventions aimed at reducing demand for anatomical parts for both ceremonial and medicinal uses.
  • Protected area expansion will also benefit this species by increasing Leopard densities in core areas and creating a more resilient population as individuals are free to roam across greater areas. The most important population is undoubtedly the Kruger National Park, and its adjacent private game reserves. The establishment of larger and better-connected protected areas, especially transfrontier conservation areas, will enhance metapopulation persistence, as had been modelled for Leopard in the Maputaland–Pondoland–Albany biodiversity hotspot (Di Minin et al. 2013). However, due to high levels of persecution and “edge effects”, even if new protected areas are created, success will largely depend on the attitudes and densities of local people (Thorn et al. 2012; Balme et al. 2010b).
  • Reintroduction is not recommended as a conservation tool at this stage. While there have been improvements in reintroduction success (Hayward et al. 2007), general consensus seems that translocation is of limited use in Leopard conservation (Athreya et al. 2011; Weilenmann et al. 2011). Furthermore, recent research on dispersal patterns in Leopards has demonstrated that competition for mates was the main driver for dispersal and thus dispersal increased with local subpopulation density (Fattebert et al. 2015). Therefore, interventions that increase local abundance of Leopards will also restore dispersal patterns disrupted by unsustainable harvesting and thus improve connectivity between subpopulations (Fattebert et al. 2015). Interventions that reduce the unsustainable harvesting and persecution of Leopards may also thus galvanise natural recolonisation and population dynamics.
Recommendations for land managers and practitioners:
  • Monitoring frameworks should be established to track provincial population trends, facilitating effective adaptive management. This could be a combination of both intensive (annual camera trap surveys in strategic sites) and extensive (change in harvest composition, hunting success and Leopard occupancy determined by trophy photographs, hunt return forms and province-wide questionnaire surveys gauging presence/absence of Leopards).
  • Adoption of sustainable science-based hunting regulations at a national level. For example, recommend a ban on the hunting of female Leopards (Daly et al. 2005; Balme et al. 2010a), for hunting effort to be equitably distributed across Leopard range (Balme et al. 2010a), and that hunting be limited to male Leopards > 7 years old (Packer et al. 2009; Balme et al. 2012). Professional Hunters should be scrutinised for experience and efficacy in terms of identification of the right animal to be hunted. It should be a highly specialised hunt, with proper assessments done beforehand.
  • Similarly, sustainable DCA protocols for putative problem Leopards should be adopted at a national level, where there is improved record keeping of trophy hunting and DCA permits. Integrated conservation plans are necessary. For example, in response to a fragmented (Swanepoel et al. 2013), and declining Leopard population (Power 2014), North West conservation authorities have embarked upon operations, combining law enforcement and problem animal control, to restore the species population status by reintroducing injured or problem individual Leopards. Call centres should be established to assist landowners with conflict management.
  • Public awareness and education programmes should be used to encourage livestock and wildlife owners to adopt non-lethal conflict mitigation approaches to reduce the risk of depredation (Ogada et al. 2003; Rust et al. 2013; McManus et al. 2015b), or to encourage product substitution among the Shembe (and other traditional users of Leopard skins).
  • Increased enforcement of the illegal persecution or use of Leopard skins for cultural and religious purposes should be promoted.
Research priorities: Leopard research in South Africa is biased toward protected areas, and thus quantitative assessment of Leopard viability in the assessment region is hampered by a number of key areas of data deficiency (Swanepoel 2013; Balme et al. 2014). These include:
  • The effect of land type on Leopard population status and trends: there are limited data on population sizes and trends in non-protected areas.
  • The scale and scope of the illegal trade in Leopard skins for cultural/religious regalia: there are limited data on illegal harvest and persecution.
  • Further research and monitoring is needed on the effects of persecution and illegal harvest on population persistence, as well as the efficacy of education and awareness programmes in mitigating this threat. Greater effort will be needed to collate the number of DCA and trophy hunting permits issued.
  • Research on actual versus perceived impacts on game ranches via Leopard predation (Funston et al. 2013). Conversely, investigating the impact of commercial game ranching on Leopard population persistence.
  • The effectiveness of non-lethal mitigation approaches to reduce human–Leopard conflict.
  • Further clarification on genetic structure of the population and likely connectivity between subpopulations.
  • Relationships between Leopard landscape use and risk of road mortality.
The following research projects are currently ongoing:
  • Panthera: 1) Provincial Monitoring Frameworks – partnering with provincial and national conservation authorities to establish monitoring networks to track Leopard population trends at meaningful management scales; 2) Furs for Life – combatting the illegal trade in Leopard skins for cultural regalia through education, policy and the provision of faux Leopard furs.
  • Landmark Leopard and Predator Foundation: ecology of Leopards, remedial action for injured 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 (for example, 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 Board: Ingwe Leopard Project: Greater Lydenburg area; Kruger National western boundary carnivore monitoring, including the neighbouring rural areas; spatial ecology, habitat utilisation, population demographics and conservation of Leopards in the Loskop Dam Nature Reserve.
Encouraged citizen actions:
  • Report sightings on virtual museum platforms (for example, iSpot and MammalMAP), especially outside protected areas.
  • Lobby to insist on proper trophy hunting procedures and permits.
  • Conduct camera trapping surveys and submit data to local conservation authority.
  • Conduct snare removal on private land.
  • Pressure government authorities to pursue criminal cases involving this species.

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,@phdthesis{3847,
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 year = {2010}
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,@article{5229,
 author = {Hayward MW, Henschel P, O’brien J, Hofmeyr M, Balme G, Kerley GIH},
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,@article{15951,
 author = {Di Minin E, Hunter LT, Balme GA, Smith RJ, Goodman PS, Slotow R.},
 journal = {PloS One},
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,@article{15951,
 author = {Marker L, Dickman A, Schumann M.},
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 pages = {28--32},
 title = {Using livestock guarding dogs as a conflict resolution strategy on Namibian farms},
 volume = {8},
 year = {2005}
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,@article{15954,
 author = {Nowell, K. and Jackson, P. (compilers and editors)},
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,@techreport{15954,
 author = {Ray, J.C.; Hunter, L. & J. Zigouris},
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,@article{15954,
 author = {Swanepoel, L.H., Lindsey, P., Somers, M.J., Hoven, W.V. and Dalerum, F.},
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 title = {Extent and fragmentation of suitable leopard habitat in South Africa},
 volume = {16},
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 author = {Athreya V, Odden M, Linnell JD, Karanth KU.},
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 author = {Balme GA, Batchelor A, Woronin Britz N, Seymour G, Grover M, Hes L, Macdonald DW, Hunter LT.},
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 title = {Reproductive success of female leopards <i>Panthera pardus</i>: the importance of top-down processes},
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 author = {Balme GA, Hunter LT, Braczkowski AR.},
 journal = {PloS One},
 pages = {e35209.},
 title = {Applicability of age-based hunting regulations for African leopards.},
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 author = {Macdonald  and Loveridge },
 booktitle = {An adaptive management approach to trophy hunting of leopards <i>Panthera pardus</i>: a case study from KwaZulu-Natal, South Africa},
 editor = {Balme GA, Hunter LT, Goodman P, Ferguson H, Craigie J, Slotow R.},
 pages = {341--352},
 publisher = {Oxford University Press},
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 year = {2010}
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,@article{15954,
 author = {Balme GA, Hunter LT, Slotow R.},
 journal = {Animal Behaviour},
 pages = {589--598},
 title = {Feeding habitat selection by hunting leopards <i>Panthera pardus</i> in a woodland savanna: prey catchability versus abundance.},
 volume = {74},
 year = {2007}
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,@article{15954,
 author = {Balme GA, Lindsey PA, Swanepoel LH, Hunter LTB.},
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 author = {Balme GA, Slotow R, Hunter LT.},
 journal = {Biological Conservation},
 pages = {2681--2690},
 title = {Impact of conservation interventions on the dynamics and persistence of a persecuted leopard (<i>Panthera pardus</i>) population},
 volume = {142},
 year = {2009}
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,@article{15954,
 author = {Balme GA, Slotow R, Hunter LTB.},
 journal = {Animal Conservation},
 pages = {315--323},
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 volume = {13},
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,@article{15954,
 author = {Bothma J du P.},
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,@article{15954,
 author = {Bothma J du P, le Riche EAN.},
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,@article{15954,
 author = {Chase Grey JN, Kent VT, Hill RA.},
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 author = {Dalerum F, Shults B, Kunkel K.},
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 title = {Estimating sustainable harvest in wolverine populations using logistic regression},
 volume = {72},
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,@techreport{15954,
 address = {Bloemfontein, South Africa.},
 author = {Du Plessis H, Smit GN.},
 institution = {Department of Animal, Wildlife and Grassland Sciences, University of the Free State},
 title = {The Development and Final Testing of an Electrified Leopard Proof Game Fence on the Farm Maseque},
 year = {1999}
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,@article{15954,
 author = {Fattebert J, Balme G, Dickerson T, Slotow R, Hunter L.},
 journal = {PLoS One},
 pages = {e0122355},
 title = {Density-dependent natal dispersal patterns in a leopard population recovering from over-harvest},
 volume = {10},
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,@article{15954,
 author = {Fattebert J, Dickerson T, Balme G, Slotow R, Hunter L.},
 journal = {South African Journal of Wildlife Research},
 pages = {61--67},
 title = {Long-distance natal dispersal in leopard reveals potential for a three-country metapopulation.},
 volume = {43},
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 author = {Funston PJ, Groom RJ, Lindsey PA.},
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 volume = {8},
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,@article{15954,
 author = {Grobler JH, Wilson VJ.},
 journal = {Arnoldia},
 pages = {1--10},
 title = {Food of the leopard <i>Panthera pardus</i> (Linn.) in the Rhodes Matopos National Park, Rhodesia, as determined by faecal analysis.},
 volume = {5},
 year = {1972}
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,@article{15954,
 author = {Hayward MW, Adendorff J, Moolman L, Hayward GJ, Kerley GI.},
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 volume = {45},
 year = {2007}
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,@inbook{15954,
 address = {London, UK},
 author = {Kingdon  and Hoffmann },
 booktitle = {<i>Panthera pardus</i> Leopard},
 editor = {Hunter, L., Henschel, P, and Ray J.C.},
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 year = {2013}
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,@article{15954,
 author = {Jooste E, Pitman RT, Van Hoven W, Swanepoel LH.},
 journal = {Folia Primatologica},
 pages = {353--360},
 title = {Unusually high predation on chacma baboons (<i>Papio ursinus</i>) by female leopards (<i>Panthera pardus</i>) in the Waterberg Mountains, South Africa.},
 volume = {83},
 year = {2013}
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,@article{15954,
 author = {Jorge AA, Vanak AT, Thaker M, Begg C, Slotow R.},
 journal = {Conservation Biology},
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 title = {Costs and benefits of the presence of leopards to the sport-hunting industry and local communities in Niassa National Reserve, Mozambique.},
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 author = {Lindsey PA, Du Toit JT, Mills MGL.},
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 title = {Attitudes of ranchers towards African wild dogs <i>Lycaon pictus</i>: conservation implications on private land},
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 year = {2005}
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,@techreport{15954,
 author = {Lindsey PA, Marnewick K, Balme G, Swanepoel LH.},
 institution = {Endangered Wildlife Trust},
 title = {Non detriment finding assessment for the trophy hunting of leopards in South Africa},
 year = {2011}
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 author = {Martin RB, De Meulenaer T.},
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 volume = {5},
 year = {1986}
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,@article{15954,
 author = {Norton PM, Lawson AB, Henley SR, Avery G.},
 journal = {South African Journal of Wildlife Research},
 pages = {47--52},
 title = {Prey of leopards in four mountainous areas of the south-western Cape Province.},
 volume = {16},
 year = {1986}
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,@article{15954,
 author = {Ogada MO, Woodroffe R, Oguge NO, Frank LG.},
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 title = {Limiting depredation by African carnivores: the role of livestock husbandry.},
 volume = {17},
 year = {2003}
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,@article{15954,
 author = {Packer C, Brink H, Kissui BM, Maliti H, Kushnir H, Caro T.},
 journal = {Conservation Biology},
 pages = {142--153},
 title = {Effects of trophy hunting on lion and leopard populations in Tanzania.},
 volume = {25},
 year = {2011}
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,@article{15954,
 author = {Pienaar U de V.},
 journal = {Koedoe},
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 volume = {12},
 year = {1969}
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,@article{15954,
 author = {Pitman RT, Kilian PJ, Ramsay PM, Swanepoel LH.},
 journal = {South African Journal of Wildlife Research},
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 title = {Foraging and habitat specialization by female leopards (<i>Panthera pardus</i>) in the Waterberg Mountains of South Africa},
 volume = {43},
 year = {2013}
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,@article{15954,
 author = {Pitman RT, Swanepoel LH, Hunter L, Slotow R, Balme GA.},
 journal = {Biodiversity and Conservation},
 pages = {1975--1987},
 title = {The importance of refugia, ecological traps and scale for large carnivore management.},
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 year = {2015}
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,@techreport{15954,
 author = {Purchase G, Mateke C.},
 title = {The state of knowledge regarding leopard (<i>Panthera pardus</i>) in three range states (Mozambique, Zambia and Zimbabwe) in the context of improving management of trophy hunting.},
 year = {2008}
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,@article{15954,
 author = {Ropiquet A, Knight AT, Born C, Martins Q, Balme G, Kirkendall L, Hunter L, Senekal C, Matthee CA.},
 journal = {Comptes Rendus Biologies},
 pages = {728--737},
 title = {Implications of spatial genetic patterns for conserving African leopards.},
 volume = {338},
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 author = {Soulé ME, Estes JA, Berger J, Del Rio CM.},
 journal = {Conservation Biology},
 pages = {1238--1250},
 title = {Ecological effectiveness: conservation goals for interactive species.},
 volume = {17},
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 author = {Spong G, Johansson M, Björklund M.},
 journal = {Molecular Ecology},
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 title = {High genetic variation in leopards indicates large and long-term stable effective population size.},
 volume = {9},
 year = {2000}
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 author = {St John FAV, Keane AM, Edwards-Jones G, Jones L, Yarnell RW, Jones JPG.},
 journal = {Proceedings of the Royal Society of London B: Biological Sciences},
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 title = {Identifying indicators of illegal behaviour: carnivore killing in human-managed landscapes.},
 volume = {279},
 year = {2011}
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,@electronic{15954,
 author = {Stein AB, Athreya V, Gerngross P, Balme G, Henschel P, Karanth U, Miquelle D, Rostro S, Kamler JF, Laguardia A.},
 publisher = {The IUCN Red List of Threatened Species 2016: e.T15954A50659089.},
 title = {<i>Panthera pardus</i>},
 year = {2016}
}
,@article{15954,
 author = {Stuart CT, Stuart TD.},
 journal = {Revue de Zoologie Africaine},
 pages = {135--137},
 title = {Prey of leopards in the western Soutpansberg, South Africa.},
 volume = {107},
 year = {1993}
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,@phdthesis{15954,
 author = {Swanepoel LH.},
 school = {University of Pretoria},
 title = {Ecology and conservation of leopards, <i>Panthera pardus</i>, on selected game ranches in the Waterberg region, Limpopo, South Africa.},
 year = {2008}
}
,@phdthesis{15954,
 author = {Swanepoel LH.},
 school = {University of Pretoria},
 title = {Viability of leopards <i>Panthera pardus</i> (Linnaeus, 1758) in South Africa.},
 year = {2013}
}
,@article{15954,
 author = {Swanepoel LH, Somers MJ, Dalerum F.},
 journal = {Wildlife Biology},
 pages = {263--268},
 title = {Density of leopards <i>Panthera pardus</i> on protected and non-protected land in the Waterberg Biosphere, South Africa},
 volume = {21},
 year = {2015}
}
,@article{15954,
 author = {Swanepoel LH, Somers MJ, van Hoven W, Schiess-Meier M, Owen C, Snyman A, Martins Q, Senekal C, Camacho G, Boshoff W, Dalerum F.},
 journal = {Oryx},
 pages = {595--603},
 title = {Survival rates and causes of mortality of leopards <i>Panthera pardus</i> in southern Africa.},
 volume = {49},
 year = {2015}
}
,@article{15954,
 author = {Swanepoel LH, Lindsey P, Somers MJ, Van Hoven W, Dalerum F.},
 journal = {South African Journal of Wildlife Research},
 pages = {115--134},
 title = {The relative importance of trophy harvest and retaliatory killing of large carnivores: South African leopards as a case study},
 volume = {44},
 year = {2014}
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,@article{15954,
 author = {Tensen L, Roelofs D, Swanepoel LH.},
 journal = {South African Journal of Wildlife Research},
 pages = {193--197},
 title = {A note on the population structure of leopards (<i>Panthera pardus</i>) in South Africa.},
 volume = {44},
 year = {2011}
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,@article{15954,
 author = {Thorn M, Green M, Scott D, Marnewick K.},
 journal = {Biodiversity and Conservation},
 pages = {1715--1730},
 title = {Characteristics and determinants of human–carnivore conflict in South African farmland.},
 volume = {22},
 year = {2013}
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,@article{15954,
 author = {Uphyrkina, O., Johnson, W.E., Quigley, H.B., Miquelle, D.G., Marker, L., Bush, M.E. and O'Brien, S.J.},
 journal = {Molecular Ecology},
 pages = {2617--2633},
 title = {Phylogenetics, genome diversity and origin of modern leopard, <i>Panthera pardus</i>},
 volume = {10},
 year = {2001}
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,@article{15954,
 author = {Weilenmann M, Gusset M, Mills DR, Gabanapelo T, Schiess-Meier M.},
 journal = {Wildlife Research},
 pages = {702--707},
 title = {Is translocation of stock-raiding leopards into a protected area with resident conspecifics an effective management tool?},
 volume = {37},
 year = {2011}
}
,]
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