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.population viability analysis | threat assessment | protected areas | landscape conservation | megafauna T he spread and dominance of humans across the world during the Anthropocene have precipitated a sixth global biodiversity extinction crisis (1). To maximize biodiversity retention through this period of rapid change, scarce conservation resources need to be targeted toward species and ecosystems that are most 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 modeling shows that, where cheetah population growth rates are suppressed outside protected areas, extinction risk increases markedly. This result can be generalized to other "protection-reliant" species, and a decision tree is provided to improve their extinction risk estimation. Ultimately, the persistence of protection-reliant species depends on their survival outside and inside protected areas and requires a holistic approach to conservation that engages rather than alienates local communities.

a b s t r a c tLarge carnivores are in rapid global decline, with a broad array of consequences for the ecosystems they inhabit. To efficiently detect and address these declines requires unbiased and precise demographic data. Unfortunately, the characteristics that make large carnivores extinction-prone also pose serious challenges to obtaining these data. Rapid survey methods exist, but provide only relative measures of abundance, cannot detect declines before they become large, and provide little or no information about the causes of decline. African lions (Panthera leo) are declining throughout their range, making accurate monitoring of remaining populations urgent. We provide statistically rigorous estimates of population size, trends, survival rate and age-sex structure from Zambia's South Luangwa lion population from 2008 to 2012, just prior to cessation of hunting in 2013. Mark-recapture models fit to data from intensive monitoring of 210 individual lions in 18 prides and 14 male coalitions indicated a declining population, low recruitment, low sub-adult and adult male survival, depletion of adult males, and a senescing adult female population. Trophy hunting was the leading cause of death, with 46 males harvested. Based on these data we recommend continuing the hunting ban at least to 2016 to allow recovery, with substantially reduced quotas, age-limits, and effective trophy monitoring mandated thereafter should hunting resume. Similar data from intensive monitoring of key Zambian lion populations is required to evaluate effects of the hunting ban and provide management guidance. Effectively integrating intensive long-term monitoring and rapid survey methods should be a priority for future management and monitoring of carnivore species.

Human activities on the periphery of protected areas can limit carnivore populations, but measurements of the strength of such effects are limited, largely due to difficulties of obtaining precise data on population density and survival. We measured how density and survival rates of a previously unstudied leopard population varied across a gradient of protection and evaluated which anthropogenic activities accounted for observed patterns. Insights into this generalist's response to human encroachment are likely to identify limiting factors for other sympatric carnivore species. Motion‐sensitive cameras were deployed systematically in adjacent, similarly sized, and ecologically similar study areas inside and outside Zambia's South Luangwa National Park (SLNP) from 2012 to 2014. The sites differed primarily in the degree of human impacts: SLNP is strictly protected, but the adjacent area was subject to human encroachment and bushmeat poaching throughout the study, and trophy hunting of leopards prior to 2012. We used photographic capture histories with robust design capture–recapture models to estimate population size and sex‐specific survival rates for the two areas. Leopard density within SLNP was 67% greater than in the adjacent area, but annual survival rates and sex ratios did not detectably differ between the sites. Prior research indicated that wire‐snare occurrence was 5.2 times greater in the areas adjacent to the park. These results suggest that the low density of leopards on the periphery of SLNP is better explained by prey depletion, rather than by direct anthropogenic mortality. Long‐term spatial data from concurrent lion studies suggested that interspecific competition did not produce the observed patterns. Large carnivore populations are often limited by human activities, but science‐based management policies depend on methods to rigorously and quantitatively assess threats to populations of concern. Using noninvasive robust design capture–recapture methods, we systematically assessed leopard density and survival across a protection gradient and identified bushmeat poaching as the likely limiting factor. This approach is of broad value to evaluate the impacts of anthropogenic activities on carnivore populations that are distributed across gradients of protection.

If access to food is affected by the risk of predation, then the number of individuals killed by predators is an incomplete measure of the limiting effect of predation. Nonetheless, it is often assumed that the costs of antipredator responses (risk effects) are either small enough to be ignored or positively correlated with direct predation, and thus unlikely to alter inferences based on predation rates. These assumptions are rarely tested. Here we studied five large carnivores and ten prey species in three Zambian ecosystems to test relationships between direct predation, antipredator vigilance and trade-offs with foraging. The presence of a predator caused vigilance to increase by a factor of 2.4, with substantial variation among prey species in the strength of this response. This was associated with a 28% decrease in the proportion of individuals foraging, a trade-off that was consistent across species. We detected no correlation between direct predation and the strength of antipredator responses, which undermines the gambit of ignoring risk effects. The strength of antipredator responses was uncorrelated with broad attributes of predators and environments, but was correlated with attributes of prey. Responses were stronger for small species and for browsers/mixed feeders relative to grazers. It has previously been noted that small ungulates face higher rates of direct predation. Building on this inference, our results suggest that carnivore loss/restoration will also have stronger behaviorally-mediated effects on small ungulates, particularly browsers and mixed feeders. If such species increase their representation where carnivores are depleted, then cascading effects on vegetation would be expected.

Factors that limit African lion populations are manifold and well-recognized, but their relative demographic effects remain poorly understood, particularly trophy hunting near protected areas. We identified and monitored 386 individual lions within and around South Luangwa National Park, Zambia, for five years (2008–2012) with trophy hunting and for three additional years (2013–2015) during a hunting moratorium. We used these data with mark-resight models to estimate the effects of hunting on lion survival, recruitment, and abundance. The best survival models, accounting for imperfect detection, revealed strong positive effects of the moratorium, with survival increasing by 17.1 and 14.0 percentage points in subadult and adult males, respectively. Smaller effects on adult female survival and positive effects on cub survival were also detected. The sex-ratio of cubs shifted from unbiased during trophy-hunting to female-biased during the moratorium. Closed mark-recapture models revealed a large increase in lion abundance during the hunting moratorium, from 116 lions in 2012 immediately preceding the moratorium to 209 lions in the last year of the moratorium. More cubs were produced each year of the moratorium than in any year with trophy hunting. Lion demographics shifted from a male-depleted population consisting mostly of adult (≥4 years) females to a younger population with more (>29%) adult males. These data show that the three-year moratorium was effective at growing the Luangwa lion population and increasing the number of adult males. The results suggest that moratoria may be an effective tool for improving the sustainability of lion trophy hunting, particularly where systematic monitoring, conservative quotas, and age-based harvesting are difficult to enforce.

Ungulate populations face declines across the globe, and populations are commonly conserved by using protected areas. However, assessing the effectiveness of protected areas in conserving ungulate populations has remained difficult. Using herd size data from four years of line transect surveys and distance sampling models, we modeled population densities of four important herbivore species across a gradient of protection on the edge of Zambia’s South Luangwa National Park (SLNP) while accounting for the role of various ecological and anthropogenic variables. Our goal was to test whether protection was responsible for density dynamics in this protection gradient, and whether a hunting moratorium impacted herbivore densities during the studies. For all four species, we estimated lower densities in partially protected buffer areas adjacent to SLNP (ranging from 4.5-fold to 13.2-fold lower) compared to protected parklands. Density trends through the study period were species-specific, with some species increasing, decreasing, or remaining stable in all or some regions of the protection gradient. Surprisingly, when controlling for other covariates, we found that these observed differences were not always detectably related to the level of protection or year. Our findings highlight the importance of accounting for variables beyond strata of interest in evaluating the effectiveness of a protected area. This study highlights the importance of comprehensively modeling ungulate population density across protection gradients, identifies lands within an important protection gradient for targeted conservation and monitoring, documents prey depletion and expands our understanding on the drivers in a critical buffer area in Zambia.

Large carnivores have experienced considerable range contraction, increasing the importance of movement across human-altered landscapes between small, isolated populations. African wild dogs (Lycaon pictus) are exceptionally wide-ranging, and recolonization is an important element of their persistence at broad scales. The competition-movement-connection hypothesis suggests that adaptations to move through areas that are unfavorable due to dominant competitors might promote the ability of subordinate competitors (like wild dogs) to move through areas that are unfavorable due to humans. Here, we used hidden Markov models to test how wild dog movements were affected by the Human Footprint Index in areas inside and outside of South Luangwa National Park. Movements were faster and more directed when outside the National Park, but slowed where the human footprint was stronger. Our results can be directly and quantitatively applied to connectivity planning, and we use them to identify ways to better understand differences between species in recent loss of connectivity.

Background Prey depletion is a threat to the world’s large carnivores, and is likely to affect subordinate competitors within the large carnivore guild disproportionately. African lions limit African wild dog populations through interference competition and intraguild predation. When lion density is reduced as a result of prey depletion, wild dogs are not competitively released, and their population density remains low. Research examining distributions has demonstrated spatial avoidance of lions by wild dogs, but the effects of lions on patterns of movement have not been tested. Movement is one of the most energetically costly activities for many species and is particularly costly for cursorial hunters like wild dogs. Therefore, testing how top-down, bottom-up, and anthropogenic variables affect movement patterns can provide insight into mechanisms that limit wild dogs (and other subordinate competitors) in resource-depleted ecosystems. Methods We measured movement rates using the motion variance from dynamic Brownian Bridge Movement Models (dBBMMs) fit to data from GPS-collared wild dogs, then used a generalized linear model to test for effects on movement of predation risk from lions, predictors of prey density, and anthropogenic and seasonal variables. Results Wild dogs proactively reduced movement in areas with high lion density, but reactively increased movement when lions were immediately nearby. Predictors of prey density had consistently weaker effects on movement than lions did, but movements were reduced in the wet season and when dependent offspring were present. Conclusion Wild dogs alter their patterns of movement in response to lions in ways that are likely to have important energetic consequences. Our results support the recent suggestion that competitive limitation of wild dogs by lions remains strong in ecosystems where lion and wild dog densities are both low as a result of anthropogenic prey depletion. Our results reinforce an emerging pattern that movements often show contrasting responses to long-term and short-term variation in predation risk.