An integrated population model reveals source-sink dynamics for competitively subordinate African wild dogs linked to anthropogenic prey depletion.

Many African large carnivore populations are declining due to decline of the herbivore populations on which they depend. The densities of apex carnivores like the lion and spotted hyena correlate strongly with prey density, but competitively subordinate carnivores like the African wild dog benefit from competitive release when the density of apex carnivores is low, so the expected effect of a simultaneous decrease in resources and dominant competitors is not obvious. Wild dogs in Zambia's South Luangwa Valley Ecosystem occupy four ecologically similar areas with well-described differences in the densities of prey and dominant competitors due to spatial variation in illegal offtake. We used long-term monitoring data to fit a Bayesian integrated population model (IPM) of the demography and dynamics of wild dogs in these four regions. The IPM used Leslie projection to link a Cormack-Jolly-Seber model of area-specific survival (allowing for individual heterogeneity in detection), a zero-inflated Poisson model of area-specific fecundity and a state-space model of population size that used estimates from a closed mark-capture model as the counts from which (latent) population size was estimated. The IPM showed that both survival and reproduction were lowest in the region with the lowest density of preferred prey (puku, Kobus vardonii and impala, Aepyceros melampus), despite little use of this area by lions. Survival and reproduction were highest in the region with the highest prey density and intermediate in the two regions with intermediate prey density. The population growth rate ( λ ) was positive for the population as a whole, strongly positive in the region with the highest prey density and strongly negative in the region with the lowest prey density. It has long been thought that the benefits of competitive release protect African wild dogs from the costs of low prey density. Our results show that the costs of prey depletion overwhelm the benefits of competitive release and cause local population decline where anthropogenic prey depletion is strong. Because competition is important in many guilds and humans are affecting resources of many types, it is likely that similarly fundamental shifts in population limitation are arising in many systems.

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.

Long-term data reveal fitness costs of anthropogenic prey depletion for a subordinate competitor, the African wild dog (Lycaon pictus).

Within carnivore guilds, dominant competitors (e.g., lions, Panthera leo) are limited primarily by the density of prey, while subordinate competitors (e.g., African wild dogs, Lycaon pictus) have been limited by the density of dominant competitors. Historically, the fitness and population density of subordinate competitors have not been tightly linked to prey density. However, populations of large herbivores have declined substantially across sub-Saharan Africa due to human impacts, and where prey depletion is severe, fitness costs for competitive subordinates may begin to outweigh the benefits of competitive release. Using long-term intensive monitoring of African wild dogs in Zambia's Luangwa Valley Ecosystem (LVE), we tested the effects of prey depletion on survival and reproduction. We hypothesized that African wild dog fitness would be lower in prey-depleted areas, despite lower lion densities. Our study area included four contiguous regions that varied in protection level, prey density, and lion density. We fit Bayesian Cormack-Jolly-Seber and closed-capture models to estimate effects on survival and population density, and generalized linear models to estimate effects on reproductive success. We found that the LVE is a stronghold for African wild dogs, with an estimated median density of 4.0 individuals/100 km2. Despite this high density, survival and reproduction differed among regions, and both components of fitness were substantially reduced in the region with the lowest prey density. Anthropogenic prey depletion is becoming an important limiting factor for African wild dogs. If prey depletion (or any other form of habitat degradation) becomes severe enough that its fitness costs outweigh the benefits of competitive release, such changes can fundamentally alter the balance between limiting factors for competitively subordinate species.

African wild dog movements show contrasting responses to long and short term risk of encountering lions: analysis using dynamic Brownian bridge movement models.

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.

Hidden Markov Models reveal a clear human footprint on the movements of highly mobile African wild dogs.

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.

Do protection gradients explain patterns in herbivore densities? An example with ungulates in Zambia's Luangwa Valley.

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.

Quantifying lion (Panthera leo) demographic response following a three-year moratorium on trophy hunting.

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.

Effects of a protection gradient on carnivore density and survival: an example with leopards in the Luangwa valley, Zambia.

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.