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Disappearing spots: The global decline of cheetah andwhat it means for conservationSarah M. Durant[1],[2],[3]*, Nicholas Mitchell1,2, Rosemary Groom1,2, Nathalie Pettorelli1,3, Audrey Ipavec1,2, Andrew PJacobson1,[4], Rosie Woodroffe1,3, Monika Böhm1,3, Luke T. B. Hunter[5], Matt Becker[6],[7], Femke Broekuis[8],[9], SultanaBashir1, Leah Andresen[10], Ortwin Aschenborn[11], Mohammed Beddiaf[12], Farid Belbachir[13], Amel Belbachir-Bazi13,Ali Berbash[14], Iracelma Brandao de Matos Machado[15], Christine Breitenmoser[16],[17], Monica Chege[18], DeonCilliers[19], Harriet Davies-Mostert[20], Amy J Dickman8, Ezequiel Fabiano[21], Mohammad Farhadinia8, Paul Funston5,Philipp Henschel5, Jane Horgan[22], Hans de Iongh[23], Houman Jowkar[24],[25], Rebecca Klein23, Peter A. Lindsey5,Laurie Marker[26], Kelly Marnewick20, Joerg Melzheimer[27], Johnathan Merkle6, Jassiel Msoka[28], Maurus Msuha[29],Helen O'Neill1,3, Megan Parker[30], Gianetta Purchase1, Sahailou Samaila[31], Yohanna Saidu[32], Abdoulkarim Samna30,Anne Schmidt-Küentzel25, Eda Selebatso[33], Etotépé A. Sogbohossou[34], Alaaeldin Soultan[35], Emma Stone[36],Esther van der Meer[37], Rudie van Vuuren[38], Mary Wykstra[39], Kim Young-Overton5

[1] Institute of Zoology, Zoological Society of London, London, UK [2] Wildlife Conservation Society, New York, USA [3] Department of Genetics, Evolutionand Environment, University College London, London, UK [4] Department of Geography, University College London, London, UK [5] Panthera, New York,USA [6] Zambia Carnivore Programme, Mfuwe, Zambia [7] Conservation Biology and Ecology Program, Department of Ecology, Montana State University,USA [8] Wildlife Conservation Research Unit, Department of Zoology, University of Oxford, UK [9] Mara Cheetah Project, Kenya Wildlife Trust, Kenya [10]Nelson Mandela Metropolitan University, South Africa [11] Ministry of Environment and Tourism, Namibia [12] Office National du Parc Culturel du TassiliN'ajjer, Algeria [13] Université de Béjaïa, Algeria [14] Environment General Authority, Libya [15] Ministry of Agriculture, Angola [16] KORA, Switzerland [17]IUCN/SSC Cat Specialist Group, Switzerland [18] Kenya Wildlife Service, Kenya [19] Cheetah Outreach, South Africa [20] Endangered Wildlife Trust, SouthAfrica [21] University of Namibia, Namibia [22] Cheetah Conservation Botswana, Botswana [23] Institute of Environmental Sciences, Leiden University,Netherlands [24] Persian Wildlife and Heritage Foundation, Iran [25]Conservation of Asiatic Cheetah Program, Iran Department of Environment,Iran [26]Cheetah Conservation Fund, Namibia [27] Leibniz Institute for Zoo and Wildlife Research, Germany [28] Department of National Parks and Wildlife, Zambia[29] Tanzania Wildlife Research Institute, Tanzania [30] Working Dogs for Conservation, USA [31] Direction de la Faune, de la Chasse et des Aires Protégée,Niger [32] Nigeria National Park Service, Nigeria [33] Consultant, Botswana [34] University of Abomey-Calavi, Benin [35] Egyptian Environmental AffairsAgency, Egypt [36] University of Bristol, UK [37] Cheetah Conservation Project, Zimbabwe [38] Naankuse Foundation, Namibia [39] Action for Cheetahs inKenya, Kenya

Submitted to Proceedings of the National Academy of Sciences of the United States of America

Establishing and maintaining protected areas (PAs) is a key tool forbiodiversity conservation. However, this approach is insufficientfor many species, particularly those that are wide-ranging andsparse. The cheetah Acinonyx jubatus exemplifies such a speciesand faces extreme challenges to its survival. Here we show that theglobal population is estimated at approximately 7,100 individualsand confined to 9% of its historical distributional range. Yet themajority of current range (77%) occurs outside of PAs, wherethe species faces multiple threats. Scenario modelling shows that,where growth rates are suppressed outside PAs, extinction ratesincrease rapidly as the proportion of population protected de-clines. Sensitivity analysis shows that growth rates within PAshave to be high if they are to compensate for declines outside.Susceptibility of cheetah to rapid decline is evidenced by recentrapid contraction in range, supporting an up-listing of IUCN RedList threat assessment to Endangered. Our results are applicableto other protection-reliant species, which may be subject to sys-tematic under-estimation of threat when there is insufficient infor-mation outside PAs. Ultimately, conserving many of these speciesnecessitates a paradigm shift in conservation towards a holisticapproach that incentivises protection and promotes sustainablehuman-wildlife coexistence across large multiple-use landscapes.

Population viability analysis | Threat assessment | Protected areas |landscape conservation | megafauna

IntroductionThe spread and dominance of humans across the world during theAnthropocene has precipitated a sixth global biodiversity extinc-tion crisis (1). In order tomaximise biodiversity retention throughthis period of rapid change, scarce conservation resources needto be targeted towards species and ecosystems that are mostthreatened. However, in the absence of complete information,reliable assessment of threat is challenging. The InternationalUnion for the Conservation of Nature (IUCN) Red List criteriaare the primary tool for identifying and categorising species-based extinction risk, enabling prioritisation of species facing

most threat (2). Yet, much of the information used for assessmentcomes from relatively well-monitored populations, usually withinPAs (3) , even though, across a species’ distributional range,populations are likely to be exposed to variable threat levels anddiffering management regimes (4).

Inaccuracies in threat assessment are particularly problem-atic for large terrestrial mammals which can be especially vulner-able to anthropogenic impacts such as habitat loss and fragmen-

Significance

Here we compile and present the most comprehensive dataavailable on cheetah distribution and status. Our analysisdemonstrates dramatic recent declines of cheetah across itsdistributional range. Most cheetah occur outside protectedareas, where they are exposed to multiple threats, yet wherethere is little information on population status. Simulationmodelling shows that where cheetah population growth ratesare suppressed outside protected areas, extinction risk in-creases markedly. This result can be generalised to other'protection-reliant' species, and a decision tree is provided toimprove their extinction risk estimation. Ultimately, the persis-tence of protection-reliant species depends on their survivaloutside, and inside, protected areas, and requires a holisticapproach to conservation that engages, rather than alienates,local communities.

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Fig. 1. Known cheetah distribution in a) Africa andb) Asia. Grey shading denotes historical range, andred shading range where cheetah are known to beresident, boundaries of PAs under IUCN categories I-IV are marked in blue.

tation, human wildlife conflict, illegal wildlife trade, and over-harvesting for bushmeat or traditional use (5-7). These threats areusually higher outside PAs, leading to systematic spatial variationin population status according to levels of protection. Yet, thisspatial variation may go undetected if information on populationstatus and trends is biased towards relatively high density popu-lations, often found within PAs (3). Such biases are widespread,as wildlife management authorities may be required to monitorwildlife within PAs but not outside, and monitoring is usuallymore challenging outside PAs because wildlife are more elusiveand occur there at lower densities (8, 9). This leads to a lack ofinformation on populations outside PAs where they are generallymore threatened, resulting in an overly favourable assessment ofstatus.

ResultsCheetah status and threat assessment

The cheetah Acinonyx jubatus is a large carnivore that facesparticularly acute challenges during the Anthropocene. It isone of the most wide-ranging carnivores, with home rangesdocumented in excess of 3,000km2 (10, 11) and movementsof translocated animals exceeding 1,000km (11). Yet densitiesseldom exceed 0.02/km2, and have been recorded as low as0.0002/km2 (12).

Historically widespread across Africa and southwestern Asia,cheetah are now known to occur in only 9% of their past distri-butional range (Fig. 1). Not only has there been a worrying con-traction in global cheetah range, but current range is extremelyfragmented. The global population is tentatively estimated ataround 7,100 adult and adolescent cheetah distributed across 33populations (Table 1). More than half of the world’s cheetahoccur in a single transboundary population stretching across sixcountries in southern Africa (Table 1). Only one other populationcomprises more than 1,000 individuals, and most populations(91%) comprise 200 individuals or fewer. Six populations do noteven reach double digits. Ongoing population trends are largelyunknown, however of the 18 populations where trends could beassigned, 14 were judged to be in decline, three stable and onlyone stable or increasing (Table 1).

In Asia, the decline of cheetah has been particularly precipi-tous. Cheetah have been extirpated from 98% of their historicalrange, and a critically endangered population of Asiatic cheetahAcinonyx jubatus venaticus survives only in Iran (Table 1). Thisremnant population is tentatively estimated to comprise fewerthan 50 individuals distributed across three core areas of range(13). The rest of the world’s cheetah occur inAfrica, spread across

30 fragmented populations that are now restricted to only 13% oftheir historical distributional range (14-16) (Table 1; Fig. 1).

Across their surviving range, cheetah populations vary in thelevel of threat they experience. Most resident range (77%) is onunprotected landwhich supports an estimated 67%of the cheetahpopulation (Table 1). Here cheetah face increased pressures fromwidespread human-wildlife conflict; prey loss due to overhuntingand bushmeat harvesting; habitat loss and fragmentation; and il-legal trade (14-16). The species thus faces spatially heterogeneousthreats that are higher outside than inside PAs, while much of thedata available for threat assessment comes fromwithin PAs whichsupport the highest reported densities of cheetah (c. 0.02/km2)(17, 18). Populations on unprotected lands and in small or poorlymanaged PAs, where they are exposed to multiple threats, arelikely to be in decline. However, because of the considerablesurvey and monitoring effort required, particularly for a wide-ranging and elusive species like a cheetah, such declines are likelyto go undetected.

Protection and extinction riskSpatial variation in threat across protection gradients in a

species’ range is expected to affect overall extinction risk. Toassess these impacts for cheetah we used scenariomodelling to: a)explore the relationship between extinction risk and populationsize while varying both the proportion of land protected andthe growth rate on unprotected lands; and b) predict populationtrends. We assumed populations were stable when protected,as observed in large PAs (19). Our model revealed markedlyhigher extinction probabilities when the percentage of land underprotection was low and when growth rates outside PAs wereless than replacement (Fig. 2). When there was no migrationor medium migration (5% of the subpopulation per annum)between protected and unprotected land there was a rapid in-crease in extinction rate when the proportion of land protecteddropped below 40% (Figs 2a and b). When the migration ratewas high (10% of the subpopulation per annum) extinction ratewas high even when 80% of the population was protected and thereduction in growth rate outside PAs was modest (Fig. 2c). Long-term studies of cheetah suggest migration rates of between 5%and 10% are likely to be realistic (see materials and methods).

We simulated the global cheetah population by setting theinitial population equal to the estimated population of 7,000individuals, of which 33% occurs in PAs (Table 1). When thepopulation growth rate outside PAs was 10% less than replace-ment andmigration rate was 5%of the subpopulation per annum,simulated populations declined by 53% over 15 years (Fig. 3a).When the growth rate outside PAs was 20% less than replace-

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Table 1. Regional summary of known cheetah distributional range and populations. Historical distributional range for cheetah totals33,056,767km2, comprising 23,340,522km2 African and 9,716,245km2 Asian range (see Fig. 1).

Area Name Countries Residentrange km2

Populationsize

Overallincrease↑ [1]/sta-ble↔/decrease↓

Residentrange inPAs

%rangein PAs

Populationsize in PAs

%populationin PAs

AfricaSouthern Africa 6country polygon

Angola/Botswana/M-ozambique/Namibia/SouthAfrica/Zambia

1,212,179 4,021 ↓ 283,851 23.4 1,041 25.9

Moxico Angola 25,717 26 ? 0 0.0 0 0.0Pandmatenga/H-wange/VictoriaFalls

Botswana/Zimbabwe 25,926 50 ↓ 15,551 60.0 29 58.0

Banhine Mozambique 7,266 10 ? 0 0.0 0 0.0Malilangwe/Save/G-onarezhou

Mozambique/Zimbabwe 9,922 46 ↔ 4,757 47.9 19 41.3

Kafue Zambia 26,222 65 ? 22,185 84.6 55 84.6Liuwa Zambia 3,170 20 ↑or ↔ 2,921 92.1 18 90.0Bubyana-Nuanetsi-BubyeConservancies

Zimbabwe 8,816 40 ↓ 0 0.0 0 0.0

Zambezi valley Zimbabwe 3,612 12 ↓ 2,102 58.2 7 58.3Matusadona Zimbabwe 1,422 3 ↓ 1,422 100.0 3 100.0Midlands RhinoConservancy

Zimbabwe 318 4 ↓ 0 0.0 0 0.0

Subtotal southernAfrica

1,324,570 4,297 332,789 25.1 1,172 27.3

Afar Ethiopia 4,480 11 ↓ 1,092 24.4 3 27.3Blen-Afar Ethiopia 8,170 20 ↓ 1,856 22.7 5 25.0Ogaden Ethiopia 12,605 32 ↓ 0 0.0 0 0.0Yangudi Rassa Ethiopia 3,046 8 ↓ 3,046 100.0 8 100.0Kenya/Ethiopia/SouthSudan

Ethiopia/Kenya/SouthSudan

191,180 191 ? 37,953 19.9 38 19.9

South Turkana Kenya 3,580 36 ? 1,117 31.2 11 30.6Kidepo/S SouthSudan/NW Kenya

Kenya/SouthSudan/Uganda

6,694 19 ? 1,422 21.2 4 21.1

Serengeti/Mara/Tsavo/Laikipia/Samburu

Kenya/Tanzania 280,114 1,362 ↓ 49,705 17.7 664 48.8

Badingilo NP South Sudan 8,517 85 ? 4,741 55.7 47 55.3Radom NP South Sudan 6,821 68 ? 0 0.0 0 0.0Southern NP South Sudan 14,680 147 ? 10,863 74.0 109 74.1Ruaha ecosystem Tanzania 30,820 200 ↔ 25,551 82.9 166 83.0Maasai Steppe Tanzania 20,409 51 ↓ 3,755 18.4 9 17.6Katavi-Ugalla Tanzania 23,955 60 ? 10,475 43.7 26 43.3Subtotal easternAfrica

615,071 2,290 151,576 24.6 1,090 47.6

Adrar desIfoghas/Ahaggar/Ajjer& Mali

Algeria/Mali 762,871 191 ? 98,867 13.0 25 13.0

WAP Benin/ BurkinaFaso/Niger

25,345 25 ? 20,923 82.6 21 82.6

CAR/Chad CAR/Chad 238,234 238 ? 44,396 18.6 44 18.6Termit Massif Niger 2,820 1 ? 2,820 100.0 1 100.0Air-T Niger 8,052 2 ? 8,052 100.0 2 100.0Subtotal western,central and northernAfrica

1,037,322 457 175,058 16.9 93 20.3

Total African 2,976,963 7,044 659,423 22.2 2,355 33.4AsiaCentral and EasternLandscapes

Iran 107,566 20 ↔ 41158 38.3 N/A N/A

Northern Landscape Iran 33,445 22 ↓ 18077 54.04 N/A N/AKavir Iran 5,856 1 ↓ 5,856 100 N/A N/A

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Area Name Countries Residentrange km2

Populationsize

Overallincrease↑ [1]/sta-ble↔/decrease↓

Residentrange inPAs

%rangein PAs

Populationsize in PAs

%populationin PAs

Total Asia 146,867 43 65,091 44.3 N/A N/ATotal global 3,123,830 7,087 724,514 23.2 2,355[2] 33.42

[1] Estimates of trend apply to entire polygon thus, for example, populations may increase at specific sites, even though there is anoverall decrease across the polygon. [2] Does not include Iranian cheetah[1] Estimates of trend apply to entire polygon thus, for example, populations may increase at specific sites, even though there is anoverall decrease across the polygon.[2] Does not include Iranian cheetah

Fig. 2. Scenario modelling of a population of cheetah living on unprotectedand protected lands. Starting population is 200 individuals distributed ata varying proportion between protected and unprotected lands (x axis).Multiplicative growth rate (lambda) inside PAs is 1.0 but outside PAs isallowed to vary from this rate down to 0.8. Graphs show estimated extinctionrates under three migration scenarios: a) no migration between protectedand unprotected lands; b) medium migration rate between protected andunprotected lands of 0.05 and standard deviation 0.025; and c) high migra-tion rate of 0.1 and standard deviation 0.05. Results are reported from 1,000simulations over 50 years.

ment, then the decline was 70%. Changing the migration ratehad little effect on overall population decline (see Fig. S1). Ifthe growth rate inside PAs is above replacement, then this slowsthe rate of decline, however, growth rates need to be high tocompletely mitigate against declines (Fig. 3b).

Evidence of recent cheetah population declines is consistentwithmodelling results. For example, in Zimbabwe, where cheetahdistribution is relatively well known, cheetah were distributedacross a contiguous population encompassing 132,931km2 in2007, which contracted to a fragmented population occupyingonly 49,124km2 by 2015 (16, 20, 21). This 63% range contractionover a short period and equivalent to a loss of 11% of distribu-tional range per year, was largely due to the disappearance of

Fig. 3. Simulated a) population trajectories over three generations (15 years)of the global cheetah population and b) sensitivity analysis to changes in thegrowth rate within PAs. Starting population was the current total estimatedglobal population size of 7,000 individuals with 33% of the population onprotected lands (Table 1). The dashed line depicts results from a multiplica-tive growth rate (lambda) of 0.9 on unprotected lands, the solid line 0.8.Migration rate was set at 0.05 with standard deviation 0.025. Results arereported from 1,000 simulations and all other parameters of the model areas described for Fig. 2. The grey dotted line depicts the 50% threshold for up-listing to Endangered status using IUCN Red List criteria A3b (a populationsize reduction of ≥50%, projected or suspected to be met within the nextthree generations, based on an index of abundance (28)).

cheetah outside PAs, associatedwithmajor changes in land tenure(22). The Zimbabwean cheetah population is also estimated tohave declined by at least 85% between 1999 and 2015 (20),equivalent to an annual decline of 13%. Similarly, there havebeen recent large-scale extinctions of cheetah across western andcentral Africa (23, 24). Ongoing rapid change is likely acrossthe African continent due to changes in land tenure (22); large-scale fencing (25); land grabs (26); and political instability (27).However, cheetah status in areas where they are most threatenedis usually uncertain, because those areas lack data. On this basis,in line with the precautionary approach and in the absence ofalternative information, our analysis suggests cheetah should beup-listed toEndangered under IUCNRedList criterionA3b (28).

Protection-reliant speciesOur model is generic, depending primarily on data on the

mean and variance of the growth rate, and demonstrates thatextinction risk can be seriously under-estimated if differences inpopulation growth rate on protected and unprotected land are nottaken into account. We assumed two panmictic subpopulations:one protected and one unprotected. In reality, populations arelikely to be much more fragmented, which increases extinctionrisk, as small isolated populations are more extinction-prone thanlarge connected ones (29). We also assumed the PA subpopula-tion was stable and hence unable to compensate for pressureson unprotected populations. This assumption may hold for many

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Fig. 4. Decision tree for threat-assessment of protection-reliant species

large mammal species. Indeed, given widespread evidence ofwildlife declines inmany PAs (30), our assumption of stabilitymayeven be overly optimistic. If populations are able to grow insidePAs, this will help mitigate against declines outside PAs, howevergrowth rates in excess of 8% per annum inside PAs are needed tocounteract a decline of more than 10% per annum outside PAs(Fig. 3b).

There is growing evidence that many populations are subjectto source-sink dynamics, whereby protected source populationsmay supplement declining sink populations (31). Our resultsshow that, when sources are unable to mitigate against declines,then there may be catastrophic consequences on populations.Populations of wide-ranging species are particularly vulnerable toedge effects on PA boundaries which will damage their capacity toact as sources and compensate for sinks outside (32). Worryingly,there is also increasing evidence for exacerbated sink effects, or‘ecological traps’, where species are attracted to sinks or ‘traps’that may be outside PAs, either because they harbour importantresources or to avoid competition or predation (33).

Accordingly, our modelling scenarios are not unrealistic andresults may be generalised to those other large mammal speciesthat are assessed to be protection-reliant. Such species may havesubstantial range outside PAs, yet are vulnerable to rapid anthro-pogenic change which results in populations outside PAs actingas sinks. Our analysis shows that assessment of threat may beunderestimated for protection-reliant species, requiring urgent re-assessment of extinction risk. We provide a decision tree to assistthis assessment process based on our simulation results, whichtakes account of the proportion of distribution or populationoutside PAs and evidence on threats (Fig. 4). The termprotection-reliant differs in important respects from the conservation de-

pendent subcategory within the lower risk category used in theIUCN Red List until 2001 (34). Conservation dependent speciesare not threatened, but might be so if conservation measuresare withdrawn. By contrast, protection-reliant species may oftenbe threatened, and additionally face elevated risks of extinctionbecause of increased pressures outside PAs where a substantialproportion of their populations persist.

Clearly, an accurate assessment of threat is a key step inidentifying those protection-reliant species that are most vulner-able to extinction; however for some species the PA systemmay be insufficient to secure long term survival. In the case ofcheetah, PAs support only an estimated 2,360 individuals, andmany protected areas are too small to sustain populations thatare viable in the long term. For such protection-reliant species, adifferent approach may be needed to halt declines outside PAsand reduce impacts of edge effects on populations inside PAs,in order to maintain connectivity and secure long term viabilityof populations across large multiple-use landscapes. While somehave advocated fencing to reduce edge effects, such interventionsare likely to have considerable negative impacts on ecosystemsand communities, while the massive areas required for a wideranging species like cheetah make the costs prohibitive (25).

Our analysis shows that growth rates within PAs have to beunrealistically high to fully compensate for declining populationsoutside PAs (Fig 3b), thus protection-reliant species are likely torespond better to an approach focused on increasing their growthrates on unprotected lands. Thus, safeguarding protection-reliantspecies like cheetah may require a paradigm shift in conser-vation, away from a primary focus on protection, towards aholistic framework that additionally incorporates incentive-basedapproaches (35). For this, new policy, management and finan-

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cial tools are needed that promote coexistence between peopleand wildlife outside and adjacent to PAs (36). This will requireconcerted action from governments and effective cross-sectoralengagement across the conservation and economic developmentcommunities. Securing sustainable solutions for wildlife and peo-ple will not be easy, particularly where threatened species mayshare their range with marginalised and vulnerable communities,and where human development challenges are substantial. How-ever, unless this is achieved, the future of wide-ranging and highlythreatened species such as cheetah is in doubt.

Materials and MethodsAssessing cheetah distribution and status

Distributional mapping of cheetah in Africa used an expert-based map-ping approach established for jaguar and tiger (37, 38) during IUCN/SSCconservation strategic planning workshops for cheetah and another similarlysparse and wide-ranging species, African wild dog lycaon pictus (14-16, 21).Additional map refinements were conducted during National ConservationAction or Management Planning Workshops and from published reports andscientific articles. Mapping in Asia was conducted by a small expert teamcomprising LH, MF and HJ using information from ongoing survey work inIran and from the IUCN Red List assessment for the Asian subspecies (13,39). Resident range was defined as land where the species was known to bestill resident as recognised by (i) regular detection of the species in an area,over a period of several years; and/or (ii) evidence of breeding. Population

size for each resident range polygon was estimated either from expertknowledge (based on surveys and monitoring) or using known densities frompopulations in comparable habitats and facing similar levels of threat (14-16, 21, 28). Trends for each polygon were assigned as increasing, decreasing,stable or unknown based on the expert judgement of those working at siteswithin polygons.

Simulation modellingPopulation simulations were conducted in R (40). Mean and standard

deviation in the multiplicative growth rate (lambda) in PAs was set at thevalues observed in the female cheetah population in the Serengeti NationalPark from 1982-2011 (19): i.e. with a mean of 1.0 and standard deviationof 0.13. These growth rate parameters implicitly include the impacts ofcompetitors (such as lion Panthera leo and spotted hyena Crocuta crocuta)on overall growth rate, as both these predators were present in this PA.Even in well managed PAs, high cub mortality due to predation may preventcheetah populations achieving lambda>1 (41). Outside PAs, mean lambdawas allowed to vary from 1 down to 0.8, with the standard deviation setto the same value within PAs (0.13). For each year, growth rate inside andoutside PAs was randomly chosen from a normal distribution.

Migration between subpopulations on protected and unprotected landswas assumed to be proportionate to each subpopulation, with a normal dis-tribution and a mean annual rate set at 0.0, 0.05 and 0.1. Standard deviationin migration rate was set at half the mean. The only data available from thelong term study population in the Serengeti National Park (42) records anadult and adolescent immigration rate of 0.07 of the total population peryear between 1991-2011, with a standard deviation of 0.035 (Table S1).

Additional details on the methods are provided in the SupplementaryMaterials.

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AcknowledgementsWe are very grateful to the Howard G. Buffett Foundation for support-

ing this work and to AZA SAFE for their support of the southern Africanreview workshop in 2015. We also thank all participants of the strategicplanning workshops and national action planning workshops for providinginformation on cheetah distribution. Karen Minkowski and Lisanne Petraccaprovided invaluable assistance with the distributional mapping.

We are very grateful to the Howard G. Buffett Foundation for supporting this work and toAZASAFE for their support of the southernAfrican review workshop in 2015.We also thankall participants of the strategic planning workshops and national action planning workshops

for providing information on cheetah distribution. Karen Minkowski and Lisanne Petraccaprovided invaluable assistance with the distributional mapping.

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