More than $1 billion needed annually to secure Africa's protected areas with lions.

Protected areas (PAs) play an important role in conserving biodiversity and providing ecosystem services, yet their effectiveness is undermined by funding shortfalls. Using lions (Panthera leo) as a proxy for PA health, we assessed available funding relative to budget requirements for PAs in Africa's savannahs. We compiled a dataset of 2015 funding for 282 state-owned PAs with lions. We applied three methods to estimate the minimum funding required for effective conservation of lions, and calculated deficits. We estimated minimum required funding as $978/km2 per year based on the cost of effectively managing lions in nine reserves by the African Parks Network; $1,271/km2 based on modeled costs of managing lions at ≥50% carrying capacity across diverse conditions in 115 PAs; and $2,030/km2 based on Packer et al.'s [Packer et al. (2013) Ecol Lett 16:635-641] cost of managing lions in 22 unfenced PAs. PAs with lions require a total of $1.2 to $2.4 billion annually, or ∼$1,000 to 2,000/km2, yet received only $381 million annually, or a median of $200/km2 Ninety-six percent of range countries had funding deficits in at least one PA, with 88 to 94% of PAs with lions funded insufficiently. In funding-deficit PAs, available funding satisfied just 10 to 20% of PA requirements on average, and deficits total $0.9 to $2.1 billion. African governments and the international community need to increase the funding available for management by three to six times if PAs are to effectively conserve lions and other species and provide vital ecological and economic benefits to neighboring communities.

The global decline of cheetah Acinonyx jubatus and what it means for conservation.

Establishing and maintaining protected areas (PAs) are key tools for biodiversity conservation. However, this approach is insufficient for many species, particularly those that are wide-ranging and sparse. The cheetah Acinonyx jubatus exemplifies such a species and faces extreme challenges to its survival. Here, we show that the global population is estimated at ∼7,100 individuals and confined to 9% of its historical distributional range. However, the majority of current range (77%) occurs outside of PAs, where the species faces multiple threats. Scenario modeling shows that, where growth rates are suppressed outside PAs, extinction rates increase rapidly as the proportion of population protected declines. Sensitivity analysis shows that growth rates within PAs have to be high if they are to compensate for declines outside. Susceptibility of cheetah to rapid decline is evidenced by recent rapid contraction in range, supporting an uplisting of the International Union for the Conservation of Nature (IUCN) Red List threat assessment to endangered. Our results are applicable to other protection-reliant species, which may be subject to systematic underestimation of threat when there is insufficient information outside PAs. Ultimately, conserving many of these species necessitates a paradigm shift in conservation toward a holistic approach that incentivizes protection and promotes sustainable human-wildlife coexistence across large multiple-use landscapes.

Genetic guidelines for translocations: Maintaining intraspecific diversity in the lion (Panthera leo).

Conservation translocations have become an important management tool, particularly for large wildlife species such as the lion (Panthera leo). When planning translocations, the genetic background of populations needs to be taken into account; failure to do so risks disrupting existing patterns of genetic variation, ultimately leading to genetic homogenization, and thereby reducing resilience and adaptability of the species. We urge wildlife managers to include knowledge of the genetic background of source/target populations, as well as species-wide patterns, in any management intervention. We present a hierarchical decision-making tool in which we list 132 lion populations/lion conservation units and provide information on genetic assignment, uncertainty and suitability for translocation for each source/target combination. By including four levels of suitability, from 'first choice' to 'no option', we provide managers with a range of options. To illustrate the extent of international trade of lions, and the potential disruption of natural patterns of intraspecific diversity, we mined the CITES Trade Database for estimated trade quantities of live individuals imported into lion range states during the past 4 decades. We identified 1056 recorded individuals with a potential risk of interbreeding with wild lions, 772 being captive-sourced. Scoring each of the records with our decision-making tool illustrates that only 7% of the translocated individuals were 'first choice' and 73% were 'no option'. We acknowledge that other, nongenetic factors are important in the decision-making process, and hence a pragmatic approach is needed. A framework in which source/target populations are scored based on suitability is not only relevant to lion, but also to other species of wildlife that are frequently translocated. We hope that the presented overview supports managers to include genetics in future management decisions and contributes towards conservation of the lion in its full diversity.

Conserving Africa's wildlife and wildlands through the COVID-19 crisis and beyond.

The SARS-CoV-2 virus and COVID-19 illness are driving a global crisis. Governments have responded by restricting human movement, which has reduced economic activity. These changes may benefit biodiversity conservation in some ways, but in Africa, we contend that the net conservation impacts of COVID-19 will be strongly negative. Here, we describe how the crisis creates a perfect storm of reduced funding, restrictions on the operations of conservation agencies, and elevated human threats to nature. We identify the immediate steps necessary to address these challenges and support ongoing conservation efforts. We then highlight systemic flaws in contemporary conservation and identify opportunities to restructure for greater resilience. Finally, we emphasize the critical importance of conserving habitat and regulating unsafe wildlife trade practices to reduce the risk of future pandemics.

Bushmeat hunting and extinction risk to the world's mammals.

Terrestrial mammals are experiencing a massive collapse in their population sizes and geographical ranges around the world, but many of the drivers, patterns and consequences of this decline remain poorly understood. Here we provide an analysis showing that bushmeat hunting for mostly food and medicinal products is driving a global crisis whereby 301 terrestrial mammal species are threatened with extinction. Nearly all of these threatened species occur in developing countries where major coexisting threats include deforestation, agricultural expansion, human encroachment and competition with livestock. The unrelenting decline of mammals suggests many vital ecological and socio-economic services that these species provide will be lost, potentially changing ecosystems irrevocably. We discuss options and current obstacles to achieving effective conservation, alongside consequences of failure to stem such anthropogenic mammalian extirpation. We propose a multi-pronged conservation strategy to help save threatened mammals from immediate extinction and avoid a collapse of food security for hundreds of millions of people.

Collapse of the world's largest herbivores.

Large wild herbivores are crucial to ecosystems and human societies. We highlight the 74 largest terrestrial herbivore species on Earth (body mass ≥100 kg), the threats they face, their important and often overlooked ecosystem effects, and the conservation efforts needed to save them and their predators from extinction. Large herbivores are generally facing dramatic population declines and range contractions, such that ~60% are threatened with extinction. Nearly all threatened species are in developing countries, where major threats include hunting, land-use change, and resource depression by livestock. Loss of large herbivores can have cascading effects on other species including large carnivores, scavengers, mesoherbivores, small mammals, and ecological processes involving vegetation, hydrology, nutrient cycling, and fire regimes. The rate of large herbivore decline suggests that ever-larger swaths of the world will soon lack many of the vital ecological services these animals provide, resulting in enormous ecological and social costs.

Underperformance of African protected area networks and the case for new conservation models: insights from Zambia.

Many African protected areas (PAs) are not functioning effectively. We reviewed the performance of Zambia's PA network and provide insights into how their effectiveness might be improved. Zambia's PAs are under-performing in ecological, economic and social terms. Reasons include: a) rapidly expanding human populations, poverty and open-access systems in Game Management Areas (GMAs) resulting in widespread bushmeat poaching and habitat encroachment; b) underfunding of the Zambia Wildlife Authority (ZAWA) resulting in inadequate law enforcement; c) reliance of ZAWA on extracting revenues from GMAs to cover operational costs which has prevented proper devolution of user-rights over wildlife to communities; d) on-going marginalization of communities from legal benefits from wildlife; e) under-development of the photo-tourism industry with the effect that earnings are limited to a fraction of the PA network; f) unfavourable terms and corruption which discourage good practice and adequate investment by hunting operators in GMAs; g) blurred responsibilities regarding anti-poaching in GMAs resulting in under-investment by all stakeholders. The combined effect of these challenges has been a major reduction in wildlife densities in most PAs and the loss of habitat in GMAs. Wildlife fares better in areas with investment from the private and/or NGO sector and where human settlement is absent. There is a need for: elevated government funding for ZAWA; greater international donor investment in protected area management; a shift in the role of ZAWA such that they focus primarily on national parks while facilitating the development of wildlife-based land uses by other stakeholders elsewhere; and new models for the functioning of GMAs based on joint-ventures between communities and the private and/or NGO sector. Such joint-ventures should provide defined communities with ownership of land, user-rights over wildlife and aim to attract long-term private/donor investment. These recommendations are relevant for many of the under-funded PAs occurring in other African countries.

Insights into the management of large carnivores for profitable wildlife-based land uses in African savannas.

Large African predators, especially lions (Panthera leo) and leopards (Panthera pardus), are financially valuable for ecotourism and trophy hunting operations on land also utilized for the production of other wildlife species for the same purpose. Predation of ungulates used for trophy hunting can create conflict with landholders and trade off thus exists between the value of lions and leopards and their impact on ungulate populations. Therefore productionist and conservation trade-offs are complexly graded and difficult to resolve. We investigated this with a risk-benefit analysis on a large private wildlife production area in Zimbabwe. Our model showed that lions result in substantial financial costs through predation on wild ungulates that may not be offset by profits from hunting them, whereas the returns from trophy hunting of leopards are projected to exceed the costs due to leopard predation. In the absence of additional income derived from photo-tourism the number of lions may need to be managed to minimize their impact. Lions drive important ecological processes, but there is a need to balance ecological and financial imperatives on wildlife ranches, community wildlife lands and other categories of multiple use land used for wildlife production. This will ensure the competitiveness of wildlife based land uses relative to alternatives. Our findings may thus be limited to conservancies, community land-use areas and commercial game ranches, which are expansive in Africa, and should not necessarily applied to areas where biodiversity conservation is the primary objective, even if hunting is allowed there.

Determinants of persistence and tolerance of carnivores on Namibian ranches: implications for conservation on Southern African private lands.

Changing land use patterns in southern Africa have potential to dramatically alter the prospects for carnivore conservation. Understanding these influences is essential for conservation planning. We interviewed 250 ranchers in Namibia to assess human tolerance towards and the distribution of large carnivores. Cheetahs (Acinonyx jubatus), leopards (Panthera pardus) and brown hyaenas (Hyaena brunnea) were widely distributed on Namibian farmlands, spotted hyaenas (Crocuta crocuta) had a narrower distribution, and wild dogs (Lycaon pictus) and lions (Panthera leo) are largely limited to areas near source populations. Farmers were most tolerant of leopards and least tolerant of lions, wild dogs and spotted hyaenas. Several factors relating to land use correlated consistently with carnivore-presence and landowner tolerance. Carnivores were more commonly present and/or tolerated where; wildlife diversity and biomass were higher; income from wildlife was higher; income from livestock was lower; livestock biomass was lower; in conservancies; game fencing was absent; and financial losses from livestock depredation were lower. Efforts to create conditions whereby the costs associated with carnivores are lowest, and which confer financial value to them are likely to be the most effective means of promoting carnivore conservation. Such conditions are achieved where land owners pool land to create conservancies where livestock are replaced with wildlife (or where livestock husbandry is improved) and where wildlife generates a significant proportion of ranch income. Additional measures, such as promoting improved livestock husbandry and educational outreach efforts may also help achieve coexistence with carnivores. Our findings provide insights into conditions more conducive to the persistence of and tolerance towards large carnivores might be increased on private (and even communal) lands in Namibia, elsewhere in southern and East Africa and other parts of the world where carnivore conservation is being attempted on private lands.

The trophy hunting of African lions: scale, current management practices and factors undermining sustainability.

The trophy hunting of lions Panthera leo is contentious due to uncertainty concerning conservation impacts and because of highly polarised opinions about the practice. African lions are hunted across at least ~558,000 km(2), which comprises 27-32% of the lion range in countries where trophy hunting of the species is permitted. Consequently, trophy hunting has potential to impart significant positive or negative impacts on lions. Several studies have demonstrated that excessive trophy harvests have driven lion population declines. There have been several attempts by protectionist non-governmental organisations to reduce or preclude trophy hunting via restrictions on the import and export of lion trophies. We document the management of lion hunting in Africa and highlight challenges which need addressing to achieve sustainability. Problems include: unscientific bases for quota setting; excessive quotas and off-takes in some countries; fixed quotas which encourage over-harvest; and lack of restrictions on the age of lions that can be hunted. Key interventions needed to make lion hunting more sustainable, include implementation of: enforced age restrictions; improved trophy monitoring; adaptive management of quotas and a minimum length of lion hunts of at least 21 days. Some range states have made important steps towards implementing such improved management and off-takes have fallen steeply in recent years. For example age restrictions have been introduced in Tanzania and in Niassa in Mozambique, and are being considered for Benin and Zimbabwe, several states have reduced quotas, and Zimbabwe is implementing trophy monitoring. However, further reforms are needed to ensure sustainability and reduce conservation problems associated with the practice while allowing retention of associated financial incentives for conservation.

The Zambian wildlife ranching industry: scale, associated benefits, and limitations affecting its development.

The number and area of wildlife ranches in Zambia increased from 30 and 1,420 km(2) in 1997 to 177 and ∼6,000 km(2) by 2012. Wild ungulate populations on wildlife ranches increased from 21,000 individuals in 1997 to ∼91,000 in 2012, while those in state protected areas declined steeply. Wildlife ranching and crocodile farming have a turnover of ∼USD15.7 million per annum, compared to USD16 million from the public game management areas which encompass an area 29 times larger. The wildlife ranching industry employs 1,200 people (excluding jobs created in support industries), with a further ∼1,000 individuals employed through crocodile farming. Wildlife ranches generate significant quantities of meat (295,000 kg/annum), of which 30,000 kg of meat accrues to local communities and 36,000 kg to staff. Projected economic returns from wildlife ranching ventures are high, with an estimated 20-year economic rate of return of 28%, indicating a strong case for government support for the sector. There is enormous scope for wildlife ranching in Zambia due to the availability of land, high diversity of wildlife and low potential for commercial livestock production. However, the Zambian wildlife ranching industry is small and following completion of field work for this study, there was evidence of a significant proportion of ranchers dropping out. The industry is performing poorly, due to inter alia: rampant commercial bushmeat poaching; failure of government to allocate outright ownership of wildlife to landowners; bureaucratic hurdles; perceived historical lack of support from the Zambia Wildlife Authority and government; a lack of a clear policy on wildlife ranching; and a ban on hunting on unfenced lands including game ranches. For the wildlife ranching industry to develop, these limitations need to be addressed decisively. These findings are likely to apply to other savanna countries with large areas of marginal land potentially suited to wildlife ranching.

The significance of African lions for the financial viability of trophy hunting and the maintenance of wild land.

Recent studies indicate that trophy hunting is impacting negatively on some lion populations, notably in Tanzania. In 2004 there was a proposal to list lions on CITES Appendix I and in 2011 animal-welfare groups petitioned the United States government to list lions as endangered under their Endangered Species Act. Such listings would likely curtail the trophy hunting of lions by limiting the import of lion trophies. Concurrent efforts are underway to encourage the European Union to ban lion trophy imports. We assessed the significance of lions to the financial viability of trophy hunting across five countries to help determine the financial impact and advisability of the proposed trade restrictions. Lion hunts attract the highest mean prices (US$24,000-US$71,000) of all trophy species. Lions generate 5-17% of gross trophy hunting income on national levels, the proportional significance highest in Mozambique, Tanzania, and Zambia. If lion hunting was effectively precluded, trophy hunting could potentially become financially unviable across at least 59,538 km(2) that could result in a concomitant loss of habitat. However, the loss of lion hunting could have other potentially broader negative impacts including reduction of competitiveness of wildlife-based land uses relative to ecologically unfavourable alternatives. Restrictions on lion hunting may also reduce tolerance for the species among communities where local people benefit from trophy hunting, and may reduce funds available for anti-poaching. If lion off-takes were reduced to recommended maximums (0.5/1000 km(2)), the loss of viability and reduction in profitability would be much lower than if lion hunting was stopped altogether (7,005 km(2)). We recommend that interventions focus on reducing off-takes to sustainable levels, implementing age-based regulations and improving governance of trophy hunting. Such measures could ensure sustainability, while retaining incentives for the conservation of lions and their habitat from hunting.