Habitat use of and threats to African large carnivores in a mixed-use landscape.

Large carnivores increasingly inhabit human-affected landscapes, which exhibit heterogeneity in biotic resources, anthropogenic pressures, and management strategies. Understanding large carnivore habitat use in these systems is critical for their conservation, as is the evaluation of competing management approaches and the impacts of significant land-use changes. We used occupancy modeling to investigate habitat use of an intact eastern African large carnivore guild across the 45,000 km2 Ruaha-Rungwa landscape in south-central Tanzania. We determined the relative impact on five large carnivore species of biotic, anthropogenic, and management factors at the scales of home range selection and short-term use within home ranges. We also specifically tested the effect of abandonment of trophy hunting areas on large carnivore occurrence. Patterns of habitat use differed among species. Lions (Panthera leo) appeared affected by top-down limitation, as their occurrence was significantly negatively associated with illegal human activity (ß = -0.63 [SE 0.28]). African wild dogs (Lycaon pictus), instead, were limited by biotic features; the species was negatively associated with riverine areas of high sympatric predator density (ß = -1.00 [SE 0.43]) and used less-productive habitats. Spotted hyaena (Crocuta crocuta) and leopard (Panthera pardus) persisted in more disturbed areas and across habitat types. Large carnivore occurrence was not affected by whether an area was used for photographic or trophy hunting tourism; regular law enforcement was instead a better predictor of occurrence. All species fared better in actively managed hunting areas than those that had been abandoned by operators. Overall, our findings highlight the divergent habitat requirements within large carnivore guilds and the importance of adopting an integrated approach to large carnivore conservation planning in modern systems. We also identified a novel threat to African conservation areas in the form of decreased management investments associated with the abandonment of trophy hunting areas.

Uso de Hábitat y Amenazas para los Grandes Carnívoros de África en un Paisaje de Uso Mixto Resumen Cada vez más, los grandes carnívoros habitan paisajes afectados por los humanos, los cuales presentan heterogeneidad en los recursos bióticos, las presiones antropogénicas y las estrategias de manejo. El conocimiento sobre cómo usan el hábitat los grandes carnívoros en estos sistemas es crucial para su conservación, como lo es la evaluación de las estrategias competitivas de manejo y los impactos de los cambios significativos en el uso de suelo. Usamos modelos de ocupación para investigar el uso de hábitat por parte de un gremio intacto de grandes carnívoros en el este de África a lo largo de los 45,000 km2 del paisaje Ruaha-Rungwa en el centro-sur de Tanzania. Determinamos el impacto relativo que tienen los factores bióticos, antropogénicos y de manejo sobre cinco especies de grandes carnívoros a escala de selección de extensión doméstica y uso a corto plazo dentro de la extensión doméstica. También analizamos específicamente el efecto que tiene el abandono de las áreas de caza de trofeos sobre la presencia de los grandes carnívoros. Los patrones de uso de hábitat difirieron entre las especies. Los leones (Panthera leo) parecieron estar afectados por la limitación de arriba-abajo ya que su presencia estuvo asociada negativamente de manera importante con la actividad humana ilegal (ß = - 0.63 [SE 0.28]). El perro salvaje africano (Lycaon pictus) estuvo limitado por los elementos bióticos; la especie estuvo asociada negativamente con las áreas de alta densidad simpátrica de depredadores (ß = - 1.00 [SE 0.43]) y utilizó los hábitats menos productivos. La hiena moteada (Crocuta crocuta) y el leopardo (Panthera pardus) persistieron en áreas más perturbadas y en todos los tipos de hábitat. La presencia de los grandes carnívoros no se vio afectada si el área se usaba para caza de trofeos o turismo fotográfico; la aplicación regular de la ley fue un mejor pronóstico de la presencia. A todas las especies les fue mejor en las áreas de caza con gestión activa que en aquellas abandonadas por los operadores. En general, nuestros descubrimientos resaltan los distintos requerimientos de hábitat dentro de los gremios de grandes carnívoros y la importancia de adoptar un enfoque integrado para la planeación de la conservación de estas especies en los sistemas modernos. También identificamos una amenaza nueva para las áreas de conservación africana a manera de inversiones de manejo disminuidas asociadas con el abandono de las áreas de caza de trofeos.

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.

No longer locally extinct? Tracing the origins of a lion (Panthera leo) living in Gabon.

Lions (Panthera leo) are of particular conservation concern due to evidence of recent, widespread population declines in what has hitherto been seen as a common species, robust to anthropogenic disturbance. Here we use non-invasive methods to recover complete mitochondrial genomes from single hair samples collected in the field in order to explore the identity of the Gabonese Plateaux Batéké lion. Comparison of the mitogenomes against a comprehensive dataset of African lion sequences that includes relevant geographically proximate lion populations from both contemporary and ancient sources, enabled us to identify the Plateaux Batéké lion as a close maternal relative to now extirpated populations found in Gabon and nearby Congo during the twentieth century, and to extant populations of Southern Africa. Our study demonstrates the relevance of ancient DNA methods to field conservation work, and the ability of trace field samples to provide copious genetic information about free-ranging animals.

Lion (Panthera leo) populations are declining rapidly across Africa, except in intensively managed areas.

We compiled all credible repeated lion surveys and present time series data for 47 lion (Panthera leo) populations. We used a Bayesian state space model to estimate growth rate-λ for each population and summed these into three regional sets to provide conservation-relevant estimates of trends since 1990. We found a striking geographical pattern: African lion populations are declining everywhere, except in four southern countries (Botswana, Namibia, South Africa, and Zimbabwe). Population models indicate a 67% chance that lions in West and Central Africa decline by one-half, while estimating a 37% chance that lions in East Africa also decline by one-half over two decades. We recommend separate regional assessments of the lion in the World Conservation Union (IUCN) Red List of Threatened Species: already recognized as critically endangered in West Africa, our analysis supports listing as regionally endangered in Central and East Africa and least concern in southern Africa. Almost all lion populations that historically exceeded ∼ 500 individuals are declining, but lion conservation is successful in southern Africa, in part because of the proliferation of reintroduced lions in small, fenced, intensively managed, and funded reserves. If management budgets for wild lands cannot keep pace with mounting levels of threat, the species may rely increasingly on these southern African areas and may no longer be a flagship species of the once vast natural ecosystems across the rest of the continent.

Spatial co-occurrence patterns of sympatric large carnivores in a multi-use African system.

Interspecific interactions can be a key driver of habitat use, and must be accounted for in conservation planning. However, spatial partitioning between African carnivores, and how this varies with scale, remains poorly understood. Furthermore, most studies have taken place within small or highly protected areas, rather than in the heterogeneous, mixed-use landscapes characteristic of much of modern Africa. Here, we provide one of the first empirical investigations into population-level competitive interactions among an African large carnivore guild. We collected detection/non-detection data for an eastern African large carnivore guild in Tanzania's Ruaha-Rungwa conservation landscape, over an area of ~45,000 km2. We then applied conditional co-occupancy models to investigate co-occurrence between lion, leopard, and African wild dog, at two biologically meaningful scales. Co-occurrence patterns of cheetah and spotted hyaena could not be modelled. After accounting for habitat and detection effects, we found some evidence of wild dog avoidance of lion at the home range scale, and strong evidence of fine-scale avoidance. We found no evidence of interspecific exclusion of leopard by lion; rather, positive associations were observed at both scales, suggesting shared habitat preferences. We found little evidence of leopard habitat use being affected by wild dog. Our findings also reveal some interspecific effects on species detection, at both scales. In most cases, habitat use was driven more strongly by other habitat effects, such as biotic resources or anthropogenic pressures, than by interspecific pressures, even where evidence of the latter was present. Overall, our results help shed light on interspecific effects within an assemblage that has rarely been examined at this scale. We also demonstrate the effectiveness of sign-based co-occurrence modelling to describe interspecific spatial patterns of sympatric large carnivores across large scales. We conclude by discussing the implications of our findings for large carnivore conservation in modern African systems.

Non-human predator interactions with wild great apes in Africa and the use of camera traps to study their dynamics.

The slow life histories of great apes (hereafter 'apes') combined with a growing inventory of predation incidents suggest that apes may be strongly affected by direct predation, as well as by predation risk. Predation risk may shape and increase behavioural flexibility by forcing individuals to adapt their behaviour to predator patterns. Forest leopards are an apex predator of primates in African rain forests and may represent a significant risk to ape populations. More field data are needed to further elucidate the behavioural modifications of apes in response to predation. We present research methods that combine the use of remote camera traps, capture-mark-recapture statistics and occupancy modelling to study predator-African ape relationships and potential antipredator behaviour through spatial variation in species co-occurrence patterns.

Trends and biases in African large carnivore population assessments: identifying priorities and opportunities from a systematic review of two decades of research.

African large carnivores have undergone significant range and population declines over recent decades. Although conservation planning and the management of threatened species requires accurate assessments of population status and monitoring of trends, there is evidence that biodiversity monitoring may not be evenly distributed or occurring where most needed. Here, we provide the first systematic review of African large carnivore population assessments published over the last two decades (2000-2020), to investigate trends in research effort and identify knowledge gaps. We used generalised linear models (GLMs) and generalised linear mixed models (GLMMs) to identify taxonomic and geographical biases, and investigated biases associated with land use type and author nationality. Research effort was significantly biased towards lion (Panthera leo) and against striped hyaena (Hyaena hyaena), despite the latter being the species with the widest continental range. African wild dog (Lycaon pictus) also exhibited a negative bias in research attention, although this was partly explained by its relatively restricted distribution. The number of country assessments for a species was significantly positively associated with its geographic range in that country. Population assessments were biased towards southern and eastern Africa, particularly South Africa and Kenya. Northern, western, and central Africa were generally under-represented. Most studies were carried out in photographic tourism protected areas under government management, while non-protected and trophy hunting areas received less attention. Outside South Africa, almost half of studies (41%) did not include authors from the study country, suggesting that significant opportunities exist for capacity building in range states. Overall, large parts of Africa remain under-represented in the literature, and opportunities exist for further research on most species and in most countries. We develop recommendations for actions aimed at overcoming the identified biases and provide researchers, practitioners, and policymakers with priorities to help inform future research and monitoring agendas.

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.

Historical mitochondrial diversity in African leopards (Panthera pardus) revealed by archival museum specimens.

Once found throughout Africa and Eurasia, the leopard (Panthera pardus) was recently uplisted from Near Threatened to Vulnerable by the International Union for the Conservation of Nature (IUCN). Historically, more than 50% of the leopard's global range occurred in continental Africa, yet sampling from this part of the species' distribution is only sparsely represented in prior studies examining patterns of genetic variation at the continental or global level. Broad sampling to determine baseline patterns of genetic variation throughout the leopard's historical distribution is important, as these measures are currently used by the IUCN to direct conservation priorities and management plans. By including data from 182 historical museum specimens, faecal samples from ongoing field surveys, and published sequences representing sub-Saharan Africa, we identify previously unrecognized genetic diversity in African leopards. Our mtDNA data indicates high levels of divergence among regional populations and strongly differentiated lineages in West Africa on par with recent studies of other large vertebrates. We provide a reference benchmark of genetic diversity in African leopards against which future monitoring can be compared. These findings emphasize the utility of historical museum collections in understanding the processes that shape present biodiversity. Additionally, we suggest future research to clarify African leopard taxonomy and to differentiate between delineated units requiring monitoring or conservation action.

Mapping black panthers: Macroecological modeling of melanism in leopards (Panthera pardus).

The geographic distribution and habitat association of most mammalian polymorphic phenotypes are still poorly known, hampering assessments of their adaptive significance. Even in the case of the black panther, an iconic melanistic variant of the leopard (Panthera pardus), no map exists describing its distribution. We constructed a large database of verified records sampled across the species' range, and used it to map the geographic occurrence of melanism. We then estimated the potential distribution of melanistic and non-melanistic leopards using niche-modeling algorithms. The overall frequency of melanism was ca. 11%, with a significantly non-random spatial distribution. Distinct habitat types presented significantly different frequencies of melanism, which increased in Asian moist forests and approached zero across most open/dry biomes. Niche modeling indicated that the potential distributions of the two phenotypes were distinct, with significant differences in habitat suitability and rejection of niche equivalency between them. We conclude that melanism in leopards is strongly affected by natural selection, likely driven by efficacy of camouflage and/or thermoregulation in different habitats, along with an effect of moisture that goes beyond its influence on vegetation type. Our results support classical hypotheses of adaptive coloration in animals (e.g. Gloger's rule), and open up new avenues for in-depth evolutionary analyses of melanism in mammals.

Leopard (Panthera pardus) status, distribution, and the research efforts across its range.

The leopard's (Panthera pardus) broad geographic range, remarkable adaptability, and secretive nature have contributed to a misconception that this species might not be severely threatened across its range. We find that not only are several subspecies and regional populations critically endangered but also the overall range loss is greater than the average for terrestrial large carnivores. To assess the leopard's status, we compile 6,000 records at 2,500 locations from over 1,300 sources on its historic (post 1750) and current distribution. We map the species across Africa and Asia, delineating areas where the species is confirmed present, is possibly present, is possibly extinct or is almost certainly extinct. The leopard now occupies 25-37% of its historic range, but this obscures important differences between subspecies. Of the nine recognized subspecies, three (P. p. pardus, fusca, and saxicolor) account for 97% of the leopard's extant range while another three (P. p. orientalis, nimr, and japonensis) have each lost as much as 98% of their historic range. Isolation, small patch sizes, and few remaining patches further threaten the six subspecies that each have less than 100,000 km(2) of extant range. Approximately 17% of extant leopard range is protected, although some endangered subspecies have far less. We found that while leopard research was increasing, research effort was primarily on the subspecies with the most remaining range whereas subspecies that are most in need of urgent attention were neglected.

Autosomal and mtDNA Markers Affirm the Distinctiveness of Lions in West and Central Africa.

The evolutionary history of a species is key for understanding the taxonomy and for the design of effective management strategies for species conservation. The knowledge about the phylogenetic position of the lion (Panthera leo) in West/Central Africa is largely based on mitochondrial markers. Previous studies using mtDNA only have shown this region to hold a distinct evolutionary lineage. In addition, anthropogenic factors have led to a strong decline in West/Central African lion numbers, thus, the conservation value of these populations is particularly high. Here, we investigate whether autosomal markers are concordant with previously described phylogeographic patterns, and confirm the unique position of the West/Central African lion. Analysis of 20 microsatellites and 1,454 bp of the mitochondrial DNA in 16 lion populations representing the entire geographic range of the species found congruence in both types of markers, identifying four clusters: 1) West/Central Africa, 2) East Africa, 3) Southern Africa and 4) India. This is not in line with the current taxonomy, as defined by the IUCN, which only recognizes an African and an Asiatic subspecies. There are no indications that genetic diversity in West/Central Africa lions is lower than in either East or Southern Africa, however, given this genetic distinction and the recent declines of lion numbers in this region, we strongly recommend prioritization of conservation projects in West/Central Africa. As the current taxonomic nomenclature does not reflect the evolutionary history of the lion, we suggest that a taxonomic revision of the lion is warranted.

The lion in West Africa is critically endangered.

The African lion has declined to <35,000 individuals occupying 25% of its historic range. The situation is most critical for the geographically isolated populations in West Africa, where the species is considered regionally endangered. Elevating their conservation significance, recent molecular studies establish the genetic distinctiveness of West and Central African lions from other extant African populations. Interventions to save West African lions are urgently required. However formulating effective conservation strategies has been hampered by a lack of data on the species' current distribution, status, and potential management deficiencies of protected areas (PAs) harboring lions. Our study synthesized available expert opinion and field data to close this knowledge gap, and formulate recommendations for the conservation of West African lions. We undertook lion surveys in 13 large (>500 km²) PAs and compiled evidence of lion presence/absence for a further eight PAs. All PAs were situated within Lion Conservation Units, geographical units designated as priority lion areas by wildlife experts at a regional lion conservation workshop in 2005. Lions were confirmed in only 4 PAs, and our results suggest that only 406 (273-605) lions remain in West Africa, representing <250 mature individuals. Confirmed lion range is estimated at 49,000 km², or 1.1% of historical range in West Africa. PAs retaining lions were larger than PAs without lions and had significantly higher management budgets. We encourage revision of lion taxonomy, to recognize the genetic distinctiveness of West African lions and highlight their potentially unique conservation value. Further, we call for listing of the lion as critically endangered in West Africa, under criterion C2a(ii) for populations with <250 mature individuals. Finally, considering the relative poverty of lion range states in West Africa, we call for urgent mobilization of investment from the international community to assist range states to increase management effectiveness of PAs retaining lions.

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.