Paul J. Funston

• Associate Professor
• Managing Director at Panthera

Lions are one of the world’s most iconic species but are threatened with extinction. Developing effective range-wide conservation plans are crucial but hampered by the relative lack of knowledge on specific threats facing each population and the socio-political context for conservation. Here, we present a range-wide examination of the relative frag...

Spatial patterns of and competition for resources by territorial carnivores are typically explained by two hypotheses: 1) the territorial defence hypothesis and 2) the searching efficiency hypothesis. According to the territorial defence hypothesis, when food resources are abundant, carnivore densities will be high and home ranges small. In additio...

My research work is on the evolutionary biology of African lions, with focus on behavioral ecology mechanisms for coexistance. Current projects include longterm ecological studies on Serengeti lions, Conflict mitigation through conservation translocation to foster coexistence at the interface and trophy harvest monitoring and evaluation of hunted p...

As the top predator in African ecosystems, lions have lost more than 90% of their historical range, and few countries possess strong evidence for stable populations. Translocations (broadly defined here as the capture and movement of lions for various management purposes) have become an increasingly popular action for this species, but the wide arr...

Coexistence with large carnivores poses challenges to human well-being, livelihoods, development, resource management, and policy. Even where people and carnivores have historically coexisted, traditional patterns of behavior toward large carnivores may be disrupted by wider processes of economic, social, political, and climate change. Conservation...

The degree of relatedness and an interacting unifying central member can exacerbate disease transmission within a social group. The severity of the disease depends on the route, dose and frequency of infection. Group-living social mammals thus have the ability to exacerbate the spread of the disease. Lions Panthera leo are social carnivores with in...

Prey biomass, diversity, and availability dictate predator abundances and niche structure. Increased prey biomass and availability predicts that two apex predators, African lions and spotted hyaenas, should increase in abundance. Although elephant and rhino carcasses generated by poachers released prey biomass not previously available, individual l...

The role of social cues in the reproduction of social mammals, particularly carnivores, has been thoroughly studied and documented in literature. However, environmental cues such as resources of water, food, and shelter have been identified to a lesser extent. Pregnant lions (Panthera leo) are notoriously secretive during the final stages of pregna...

Human disturbance can have a profound effect on the occurrence and distribution of wildlife. Such disturbance often extends into protected areas (PAs), particularly in countries that have undergone civil strife and lack the institutional capacity to effectively mitigate anthropogenic threats. We demonstrate the first application of a multi‐species...

Context Direct observations of animals are the most reliable way to define their behavioural characteristics; however, to obtain these observations is costly and often logistically challenging. GPS tracking allows finer-scale interpretation of animal responses by measuring movement patterns; however, the true behaviour of the animal during the peri...

We document and evaluate the use of metapopulation management to conserve a declining population of 217 cheetahs in 40 subpopulations. Metapopulation management resembles a natural metapopulation, but dispersal success, demographic rescue effects and genetic viability are enhanced by moving suitable individuals to selected habitat fragments. Unfort...

Significance Protected areas (PAs) are the cornerstone of conservation yet face funding inadequacies that undermine their effectiveness. Using the conservation needs of lions as a proxy for those of wildlife more generally, we compiled a dataset of funding in Africa’s PAs with lions and estimated a minimum target for conserving the species and mana...

Competition can have profound impacts on the structure and function of ecological communities. Despite this, the population-level effects of intraguild competition on large carnivores remain largely unknown, due to a paucity of long-term studies that focus simultaneously on competing species. Here, we comprehensively examine competitive interaction...

Habitat loss and fragmentation are major drivers of biodiversity loss, with large carnivores being particularly vulnerable due to their large home ranges. These species were once widespread across the African continent but over the past 100 years their ranges have decreased by more than 75% and there is continued pressure on the remaining free-roam...

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 i...

Aims Indicators of pending state-shifts carry value for policy makers. Predator–prey relations reflect key ecological processes that shape ecosystems. Variance in predator–prey relations may serve as a key indicator of future state-shifts. Methods Lion (Panthera leo) diet in the Kruger National Park was evaluated as such an indicator. Over the thr...

Significance Here, we compile and present the most comprehensive data available on cheetah distribution and status. Our analysis shows dramatic declines of cheetah across its distributional range. Most cheetah occur outside protected areas, where they are exposed to multiple threats, but there is little information on population status. Simulation...

Background The range, population size and trend of large carnivores are important parameters to assess their status globally and to plan conservation strategies. One can use linear models to assess population size and trends of large carnivores from track-based surveys on suitable substrates. The conventional approach of a linear model with interce...

Spotted hyaenas (Crocuta crocuta) experience a range of influences that affect their demography and are generally regulated by density-dependent mechanisms. Although widely distributed across sub-Saharan Africa surprisingly few spotted hyaena populations have reliable estimates of population sizes, and almost nothing is known about the trends in hy...

Trophy hunting plays a significant role in wildlife conservation in some contexts in various parts of the world. Yet excessive hunting is contributing to species declines, especially for large carnivores. Simulation models suggest that sustainable hunting of African lions may be achieved by restricting offtakes to males old enough to have reared a...

Interventionist conservation management of territorial large carnivores has increased in recent years, especially in South Africa. Understanding of spatial ecology is an important component of predator conservation and management. Spatial patterns are influenced by many, often interacting, factors making elucidation of key drivers difficult. We had...

Emerging infectious diseases of wildlife are of increasing concern to managers and conservation policy makers, but are often difficult to study and predict due to the complexity of host-disease systems and a paucity of empirical data. We demonstrate the use of an Approximate Bayesian Computation statistical framework to reconstruct the disease dyna...

In dismissing the conclusions of our paper, “Lion (Panthera leo) populations are declining rapidly across Africa, except in intensively managed areas,” Riggio et al. (1) misrepresent our treatment of specific survey sites and raise erroneous objections to our overall statistical approach.

Population fragmentation is threatening biodiversity worldwide. Species that once roamed vast areas are increasingly being conserved in small, isolated areas. Modern management approaches must adapt to ensure the continued survival and conservation value of these populations. In South Africa, a managed metapopulation approach has been adopted for s...

Trophy hunting is widely used in Africa to generate funding for wildlife areas.In 2015, a global media frenzy resulted from the illegal killing of a radio-collared lion, “Cecil,” by a trophy hunter in Zimbabwe. Trophy hunting is con-tentious and much of the media discourse is emotional and polarized, focusingon animal welfare and debating the value...

Significance At a regional scale, lion populations in West, Central, and East Africa are likely to suffer a projected 50% decline over the next two decades, whereas lion populations are only increasing in southern Africa. Many lion populations are either now gone or expected to disappear within the next few decades to the extent that the intensivel...

African lion populations have declined dramatically due to various anthropogenic factors. As a consequence, the majority of extant lion populations are of small size, further compromising their persistence. Lion populations in unfenced areas are more exposed to illegal hunting, snaring and poisoning, while managers of small fenced lions populations...

Lion (Panthera leo) populations on small reserves (<1000 km 2) require more intensive management than those in larger conservation areas. 'Surplus' animals are a major management concern, resulting from smaller inter-birth intervals, younger ages of first reproduction and higher survival rates. In the 1990s and early 2000s, this surplus was translo...

Emerging infectious diseases of wildlife are of increasing concern to managers and conservation policy makers, but are often diffi cult to study and predict due to the complexity of host-disease systems and a paucity of empirical data. We demonstrate the use of an Approximate Bayesian Computation statistical framework to reconstruct the disease dyn...

Research that yields conflicting results rightly causes controversy. Where methodological weaknesses are apparent, there is ready opportunity for discord within the scientific community, which may undermine the entire study. We use the debate about the role of dingoes C anis dingo in conservation in A ustralia as a case study for a phenomenon that...

In this study, we were interested in understanding if droughts influence the home range of predators such as lions, and if it does, in what ways the droughts influenced lions to adjust their home range, in response to prey availability. We monitored movements of ten lions fitted with GPS-GSM collars in order to analyze their home range and movement...

Population size estimates must be comparable through time to interpret trends in threatened carnivore populations. Because prey distribution and dynamics drive carnivore distribution, and sampling methods often utilise behavioural responses to attractants, habitat variability among sampling occasions may confound such estimates. We explored whether...

The South African lion (Panthera leo) population is highly fragmented. One-third of its wild lions occur in small (<1000 km2) reserves. These lions were reintroduced from other areas of the species’ historical range. Management practices on these reserves have not prioritized genetic provenance or heterozygosity. These trends potentially constrain...

The African lion Panthera leo is an iconic species but it has faced dramatic range reductions and possibly as few as 30,000 individuals remain in the wild. In the absence of detailed ground-based surveys, lion populations may be estimated using regression models based on prey biomass availability but these often overestimate lion densities as a res...

Managers of reintroduced lion (Panthera leo) populations in small reserves (<1000 km2) in South Africa are challenged by high rates of population increase and how best to control them. We combined data from 14 small, fenced reserves to evaluate growth rate parameters and compared them to those in larger and/or open reserves. Growth rates of lions i...

Lion populations have undergone a severe decline in West Africa. As baseline for conservation management, we assessed the group structure of lions in the Pendjari Biosphere Reserve in Benin. This reserve, composed of one National Park and two Hunting Zones, is part of the WAP transboundary complex of protected areas. Overall mean group size was 2.6...

Managers of African lions (Panthera leo) on reserves where they have been reintroduced increasingly face challenges associated with ecological regulation,genetic degradation and increased susceptibility to catastrophic events. The Lion Management Forum (LiMF) was formed in 2010 to define these challenges and explore possible solutions with the view...

The spatial ecology of Africa lions (Panthera leo) was studied from 2007 to 2009 in Waza National Park, Cameroon, by equipping individual lions with GPS/VHF radio-collars. Mean home range estimates using 100% minimum convex polygons (MCP) and 95% kernel-density estimation (KDE) were respectively 1015 km2 and 641 km2. The lions spent a considerable...

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...

The feeding ecology of lions (Panthera leo, Linnaeus 1758) was investigated in and around Waza National Park, northern Cameroon. Diet was determined using global positioning system (GPS) data of lion kill sites (clusters of GPS location points) collected using radio collars. Lions consumed 14 different prey species, with five species that were eith...

Conflict between humans and lions Panthera leo is a key factor driving population declines of lions in Africa, especially in communal lands and on the edges of small protected areas. We assessed this conflict in Waza National Park, Cameroon, in 2008 through an interview survey. A total of 207 resident and 174 nomadic pastoralists were interviewed....

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...

Conservationists often advocate for landscape approaches to wildlife management while others argue for physical separation between protected species and human communities, but direct empirical comparisons of these alternatives are scarce. We relate African lion population densities and population trends to contrasting management practices across 42...

We define African savannahs as being those areas that receive between 300 and 1,500 mm of rain annually. This broad definition encompasses a variety of habitats. Thus defined, savannahs comprise 13.5 million km2 and encompass most of the present range of the African lion (Panthera leo). Dense human populations and extensive conversion of land to hu...

Prey selection by lions (Panthera leo) reintroduced into small game reserves (typically <1000 km2) in South Africa was compared with results from larger (more naturally functioning) protected areas. Prey selection was used to calculate mean preference indices, and where sample size allowed, tested for significance of preference, as well as for comp...

In West and Central Africa large carnivores have become increasingly rare as a consequence of rapid habitat destruction and lack of resources for protected area management. The Bénoué Complex (23,394km2) in northern Cameroon is a regionally critical area for large mammal conservation. In the complex lions (Panthera leo), leopards (Panthera pardus)...

Close proximity between humans and large predators results in high levels of conflict. We investigated the extent of, and factors leading to, this conflict through focal group and individual interviews in all villages around Pendjari Biosphere Reserve, northern Benin. Livestock losses from 2000 to 2007 (n = 752) were reported to be mainly caused by...

Lions (Panthera leo) that kill livestock around the Kgalagadi Transfrontier Park have been persecuted for decades. The impact of this on the population ecology and long-term survival probability of this population has been unknown. This was largely due to a lack of basic population information. The results of a three-year intensive study from 1998...

Background/Question/Methods Bovine tuberculosis (BTB), an exotic invasive disease in Kruger National Park, was detected in Kruger’s lions during 1995. While BTB may impact individual animals, observational studies did not find population effects of BTB on lion abundance and demographics once the effects of prey biomass had been accounted for. Thes...

Aim In recent decades there has been a marked decline in the numbers of African lions (Panthera leo), especially in West Africa where the species is regionally endangered. Based on the climatological history of western Africa, we hypothesize that West and Central African lions have a unique evolutionary history, which is reflected by their genetic...

Lion populations in West and Central Africa are small and fragmented. In areas where park management is weak, threats will likely facilitate the extinction of the lion. Wildlife management requires knowledge of the population estimate. The population of lions in Waza National Park (Waza NP) was assessed by individual identification of members in th...

In South Africa, more than 30 small, enclosed game reserves have reintroduced lions over the last two decades, which now house more than 500 individuals. There is a high risk of inbreeding in these fragmented, fenced and isolated populations, which may be compounded by a lack of management guidelines. A population of 11 founder lions Panthera leo w...

Population size and trends of large carnivores are difficult to determine, but are often needed to inform conservation actions. Direct counts maintained over long time periods are extremely difficult to achieve. Indices of population sizes can be used to estimate large carnivore abundances, but are often case-, species- and site-specific. Here, we...

Context. Lion (Panthera leo Linnaeus, 1758) populations experience a range of ecological and human influences that affect their demography. Few lion populations have reliable estimates of population size, trends in these, or demographic profiles. Threats such as those imposed by diseases are thus hard to evaluate and respond to. Aims. To calibrate...

[Kiffner (2009)][1] questions the comparability of the Pleistocene tar seep records and those derived from predator playback experiments. While we acknowledge that they are not identical, the differences are not sufficient to overturn our inference of sociality in Rancho La Brea (RLB) Smilodon . He

The lion Panthera leo is one of the world's most charismatic carnivores and is one of Africa's key predators. Here, we used a large dataset from 357 lions comprehending 1.13 megabases of sequence data and genotypes from 22 microsatellite loci to characterize its recent evolutionary history. Patterns of molecular genetic variation in multiple matern...

A retrospective serosurvey of multi-host feline and canine viruses among carnivore species in southern Africa (n=1018) identified widespread pathogen exposure even in remote protected areas. In contrast to mortality experienced in East African predators, canine distemper virus (CDV) infection among African wild dogs (Lycaon pictus) in Botswana was...

Genetic variation of 12S–16S (mtDNA) and ADA and TF (nDNA) genes in lions. (A) Haplotypes and variable sites for the 12S–16S mtDNA region surveyed in lions (total length 1,882 bp). Position 1 corresponds to position 1441 of the domestic cat (Felis catus) mtDNA genome (GenBank U20753). The “-” represents a gap and “.” matches the nucleotide in the f...

Linearized genetic differentiation of host and viral genetic markers with geographic distance. Regression of linearized FST estimates [24] for lion (nDNA and mtDNA) and FIVPle (pol-RT) genetic data plotted both against the geographic distance (model assuming habitat to be arrayed in an infinite one-dimensional lattice; one-dimension isolation-by-di...

Phylogenetic relationships of the 12S–16S mtDNA lion haplotypes. Neighbour-joining tree of the 1,882 bp 12S–16S mtDNA sequences. Bootstrap values are placed at each branchpoint for the minimum evolution/maximum parsimony/maximum likelihood analyses, respectively (ME/MP/ML). Outgroups: Ppa – leopard, Panthera pardus; Pun – snow-leopard, Panthera unc...

Phylogenetic relationships of the FIVPle pol-RT sequences. Neighbour-joining tree of the 301 bp FIVPle pol-RT sequences. The distinct FIVPle subtypes were labelled A to F. Bootstrap (BPS) values are placed at each branchpoint (ME/MP/ML) and in parenthesis are the BPS values obtained for a tree established with 520 bp of FIVPle pol-RT sequence for a...

Primers used to amplify the mtDNA (12S–16S) and nDNA (ADA and TF) portions surveyed in this study. (0.06 MB PDF)

Taxon specific unique nDNA alleles in lion populations (FCA-microsatellites and ADA locus). (0.06 MB PDF)

Summary statistics for FIVPle data. (0.00 MB PDF)

List of the lion samples used in this study. (0.11 MB PDF)

Nested design and summary results of the nested clade phylogeographic analysis (NCPA) for lion mtDNA data. (A) Nested design of the mtDNA haplotype network used for the NCPA. (B) Summary results of the NCPA. RGF/IBD - Restricted gene flow/isolation by distance. LDC/FR – long distance colonization/fragmentation. (0.10 MB PDF)

Structure cluster assignment results of 357 lions based on nDNA (ADA, TF, and 22 microsatellites) and mtDNA markers. Burn-in and replication values set at 30,000 and 1,000,000, respectively. (0.06 MB PDF)

Gene diversity and frequency values in lion populations. (0.07 MB PDF)

Bottleneck analysis in lion populations using the standardized differences test and the stepwise mutation model (SMM). (0.05 MB PDF)

Results of the hierarchical AMOVA in lions for four different geographical scenarios. (0.06 MB PDF)

Lion population pairwise FST estimates. Below the diagonal mtDNA data (12S–16S) and above the diagonal microsatellite data (22 loci). (0.06 MB PDF)

Inferences concerning the lives of extinct animals are difficult to obtain from the fossil record. Here we present a novel approach to the study of extinct carnivores, using a comparison between fossil records (n=3324) found in Late Pleistocene tar seeps at Rancho La Brea in North America and counts (n=4491) from playback experiments used to estima...

The reproductive biology of a single pride of lions (Panthera leo) was studied on the 8500 ha Karongwe Game Reserve from 1999-2005. Over this period the pride consisted of between 4-11 lions, with a paired coalition of adult males during the first three years, and a single adult male for the next three years. We recorded shorter than normal interbi...

The reproductive biology of a pride of lions (Panthera leo) was studied on the 8500 ha Karongwe Game Reserve from 1999 to 2005. Over this period, the pride consisted of between four and 11 lions with a paired coalition of adult males during the first three years and a single adult male for the next three years. We recorded shorter than normal inter...

1. Many studies have investigated why males and females segregate spatially in sexually dimorphic species. These studies have focused primarily on temperate zone ungulates in areas lacking intact predator communities, and few have directly assessed predation rates in different social environments. 2. Data on the movement, social affiliation, mortal...

Buffalo herd movements, home range and habitat selection were studied for 14 months in the Sabi Sand Wildtuin, Transvaal, South Africa. The vegetation associated with the types of terrain was described in detail. Veld condition estimates and seasonal biomass changes were determined and used in the interpretation of habitat selection and home range...

The prey selection and predatory behaviour of a single pride of lions (Panthera leo) was studied on the 8500 ha Karongwe Game Reserve from 1999-2005. The study focused on the difference between prey selection in the first three years when a two-male coalition was present with a similar period thereafter when one of the males was removed and subadul...

In order to accurately assess the status of the cheetah Acinonyx jubatus it is necessary to obtain data on numbers and demographic trends. However, cheetahs are notoriously difficult to survey because they occur at very low population densities and are often shy and elusive. In South Africa the problem is further complicated in areas where land is...

Interventionist conservation management of territorial large carnivores has increased in recent years, especially in South Africa. Understanding of spatial ecology is an important component of predator conservation and management. Spatial patterns are influenced by many, often interacting, factors making elucidation of key drivers difficult. We had...

Here we document what we believe is an unusual incidence of leopard (Panthera pardus) cannibalism, where an adult male killed an adult female, presumably associated with a case of intraspecific food competition. Although the prey item, and adult impala (Aepyceros melampus) was a fairly large prey item we can not explain how this resource would have...

The impact of lion (Panthera leo) predation on buffalo (Syncerus caffer) and medium-sized ungulates over periods of changing environmental conditions in an area of the Kruger National Park was investigated. Lions were found to prey proportionally most heavily on the resident buffalo and wildebeest (Connochaetes taurinus) populations, and less so on...

An age-structured demographic matrix model is developed to determine the long-term viability of the Southern African Kgalagadi Transfrontier Park lion (Panthera leo) population that is subjected to persecution along the park boundaries. In doing so, maximum sustainable persecution rates are established, so that efficient conservation measures may b...

Life-history patterns in lions Panthera leo living in savanna woodlands of the Kruger National Park, South Africa, were investigated and compared with those of the Tanzanian ‘plains-like’ ecosystems (e.g. Serengeti Plains and Ngorongoro Crater). First, lower levels of mortality in the juvenile age classes were found in Kruger lions, which in turn,...