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Abstract 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.

Abstract Predators of similar size often compete over prey. In semi‐arid ecosystems where water is a limiting resource, prey availability can be affected by water distribution, which further increases resource competition and exacerbate conflict among predators. This can have implications for carnivore dietary competition. Hence, we evaluated the dynamics of food resource competition between African wild dogs and four competing predators (cheetahs, leopards, lions and spotted hyaenas) in different seasons and across areas with different waterhole densities in Hwange National Park, Zimbabwe. We used the frequency of occurrence of prey items found in predators' scats to analyse diet composition, overlap and prey preference. For most predators, kudu was most frequently consumed and preferred. Low and medium water‐dependent prey (medium and small‐sized) were mostly consumed by wild dogs, leopards and cheetahs. Wild dog diet overlap was high with all predators, particularly with hyaenas and lions. There were no seasonal differences in the predators diet. The diet overlap of wild dogs with lions was highest in the low waterhole density area, and wild dog diet composition did not differ significantly from the diet of lions and hyaenas. In the low waterhole density area, wild dogs and hyaenas broadened their niche breadth, and predators diet had a higher proportion of low water‐dependent prey. A low density of waterholes increased food resource competition. However, high density of waterholes, where there is more prey availability, can increase the aggregation and density of predators, and hence, increase the risks involved in interspecific competition on wild dogs. To reduce food resource competition on wild dogs, we propose to conserve larger‐bodied prey that are less dependent on water (e.g. kudu, reedbuck, eland and gemsbok). As the use of water pumping is common practice, we propose maintaining water management heterogeneity where prey which is less dependent on water can also thrive.

Abstract Wide-ranging carnivores experience tradeoffs between dynamic resource availabilities and heterogeneous risks from humans, with consequences for their ecological function and conservation outcomes. Yet, research investigating these tradeoffs across large carnivore distributions is rare. We assessed how resource availability and anthropogenic risks influence the strength of lion (Panthera leo) responses to disturbance using data from 31 sites across lions’ contemporary range. Lions avoided human disturbance at over two-thirds of sites, though their responses varied depending on site-level characteristics. Lions were more likely to exploit human-dominated landscapes where resources were limited, indicating that resource limitation can outweigh anthropogenic risks and might exacerbate human-carnivore conflict. Lions also avoided human impacts by increasing their nocturnal activity more often at sites with higher production of cattle. The combined effects of expanding human impacts and environmental change threaten to simultaneously downgrade the ecological function of carnivores and intensify human-carnivore conflicts, escalating extinction risks for many species.

Abstract 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 fragility of lion populations, examining socio-political factors alongside ecological ones. We found Ethiopia’s Maze National Park had the most ecologically fragile geographic population while Kavango-Zambezi was the least. At a country level, lion populations had highest ecological fragility in Cameroon and Malawi. When we examined socio-political fragility, Somalia was the most fragile lion range country, followed by South Sudan. When socio-political and ecological fragility were combined, lion populations in Maze National Park and Bush-Bush (Somalia) and more broadly, Somalian and Malawian lion populations were the most fragile. These insights should help inform more nuanced and appropriately targeted lion conservation plans.

Hunting is an increasingly contentious topic. Trophy hunting, whereby people hunt individual animals with desirable characteristics in order to keep body parts (e.g. horns, heads, hides, antlers) as mementos, is especially contested. Political pressure, often in the form of trophy import bans, is being applied in multiple nations, and campaigns to ban trophy hunting, or trophy imports, attract considerable media attention. However, trophy hunting often has conservation value, acting to protect habitat and provide income for local communities. Assuming that media coverage can influence public and political opinion, negative or simplistic media coverage of trophy hunting has the potential to cause adverse outcomes for conservation and local communities. Here, we analyse coverage of trophy hunting from July 2010 - June 2020 (five years before and five years after the death of Cecil the Lion) in the most popular UK media outlets (624 articles in total), assessing the overall sentiment of each article, and the species and countries covered. Ninety percent of all coverage occurred after the death of Cecil the lion, marking this event as a watershed moment in UK mainstream media depiction of trophy hunting. The overall sentiment of articles was largely against trophy hunting (63.1%), and this was more pronounced in tabloids (84.2%) than broadsheets (42.2%). Pro-trophy hunting articles were very uncommon overall (3.5%). Articles that described the complexity of trophy hunting decreased following Cecil and were most common in pre-Cecil broadsheets (35.7%, dropping to 30.6%) and rarest in post-Cecil tabloids (3.1%). Articles focussed mainly on charismatic but rarely hunted species including lion, elephant and rhino, with commonly hunted species (such as impala or Cape buffalo) only rarely being mentioned. When countries were mentioned, southern African nations predominated, with four nations (Zimbabwe, South Africa, Namibia and Botswana) being named in 68% of qualifying articles. We conclude that simplistic media depiction of trophy hunting has the potential to cause negative outcomes for conservation through its impact on public perception and political opinion.

Abstract Apex carnivores are wide‐ranging, low‐density, hard to detect, and declining throughout most of their range, making population monitoring both critical and challenging. Rapid and inexpensive index calibration survey (ICS) methods have been developed to monitor large African carnivores. ICS methods assume constant detection probability and a predictable relationship between the index and the actual population of interest. The precision and utility of the resulting estimates from ICS methods have been questioned. We assessed the performance of one ICS method for large carnivores—track counts—with data from two long‐term studies of African lion populations. We conducted Monte Carlo simulation of intersections between transects (road segments) and lion movement paths (from GPS collar data) at varying survey intensities. Then, using the track count method we estimated population size and its confidence limits. We found that estimates either overstate precision or are too imprecise to be meaningful. Overstated precision stemmed from discarding the variance from population estimates when developing the method and from treating the conversion from tracks counts to population density as a back‐transformation, rather than applying the equation for the variance of a linear function. To effectively assess the status of species, the IUCN has set guidelines, and these should be integrated in survey designs. We propose reporting the half relative confidence interval width (HRCIW) as an easily calculable and interpretable measure of precision. We show that track counts do not adhere to IUCN criteria, and we argue that ICS methods for wide‐ranging low‐density species are unlikely to meet those criteria. Established, intensive methods lead to precise estimates, but some new approaches, like short, intensive, (spatial) capture–mark–recapture (CMR/SECR) studies, aided by camera trapping and/or genetic identification of individuals, hold promise. A handbook of best practices in monitoring populations of apex carnivores is strongly recommended.

Abstract Within a large carnivore guild, subordinate competitors (African wild dog, Lycaon pictus, and cheetah, Acinonyx jubatus) might reduce the limiting effects of dominant competitors (lion, Panthera leo, and spotted hyena, Crocuta crocuta) by avoiding them in space, in time, or through patterns of prey selection. Understanding how these competitors cope with one other can inform strategies for their conservation. We tested how mechanisms of niche partitioning promote coexistence by quantifying patterns of prey selection and the use of space and time by all members of the large carnivore guild within Liuwa Plain National Park in western Zambia. Lions and hyenas specialized on wildebeest, whereas wild dogs and cheetahs selected broader diets including smaller and less abundant prey. Spatially, cheetahs showed no detectable avoidance of areas heavily used by dominant competitors, but wild dogs avoided areas heavily used by lions. Temporally, the proportion of kills by lions and hyenas did not detectably differ across four time periods (day, crepuscular, early night, and late night), but wild dogs and especially cheetahs concentrated on time windows that avoided nighttime hunting by lions and hyenas. Our results provide new insight into the conditions under which partitioning may not allow for coexistence for one subordinate species, the African wild dog, while it does for cheetah. Because of differences in responses to dominant competitors, African wild dogs may be more prone to competitive exclusion (local extirpation), particularly in open, uniform ecosystems with simple (often wildebeest dominated) prey communities, where spatial avoidance is difficult.

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.

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.

ABSTRACT Prey depletion threatens many carnivore species across the world and can especially threaten low‐density subordinate competitors, particularly if subordinates are limited to low densities by their dominant competitors. Understanding the mechanisms that drive responses of carnivore density to prey depletion is not only crucial for conservation but also elucidates the balance between top‐down and bottom‐up limitations within the large carnivore guild. To avoid predation, competitively subordinate African wild dogs typically avoid their dominant competitors (lions) and the prey rich areas they are associated with, but no prior research has tested whether this pattern persists in ecosystems with anthropogenically‐reduced prey density, and reduced lion density as a result. We used spatial data from wild dogs and lions in the prey‐depleted Greater Kafue Ecosystem to test if wild dogs continue to avoid lions (despite their low density), and consequently avoid habitats with higher densities of their dominant prey species. We found that although lion density is 3X lower than comparable ecosystems, wild dogs continue to strongly avoid lions, and consequently avoid habitats associated with their two most important prey species. Although the density of lions in the GKE is low due to prey depletion, their competitive effects on wild dogs remain strong. These effects are likely compounded by prey‐base homogenization, as lions in the GKE now rely heavily on the same prey preferred by wild dogs. These results suggest that a reduction in lion density does not necessarily reduce competition, and helps explain why wild dogs decline in parallel with their dominant competitors in ecosystems suffering from anthropogenic prey depletion. Protecting prey populations within the few remaining strongholds for wild dogs is vitally important to avoid substantial population declines. Globally, understanding the impacts of prey depletion on carnivore guild dynamics should be an increasingly important area of focus for conservation.

While telemetry technology has undoubtedly revolutionized ecological research, the impacts to conservation remain in question. Conservation is, after all, an applied discipline with research that is intentionally designed to inform policies and practices that can demonstrably protect biodiversity. Though telemetry is a tool that is commonly used to raise conservation funding, the technology itself cannot generate these policies and practices. Rather, it is the outputs of the analytical processes interrogating the data deriving from telemetry systems that can do so. This distinction is not semantic but rather fundamental to creating more actionable research. We developed conceptual frameworks to delineate the pathways by which telemetry research can be structured to inform conservation policies, practices, and the decisions of funders motivated to support conservation. We demonstrate how the application of these frameworks can reduce the research-implementation gap so as to make biodiversity conservation more effective. While our assessment uses collaring as a case study, our conceptual frameworks are applicable to all research using animal-borne technology seeking to promote the recovery of species of conservation concern.

Abstract Large herbivore migrations are imperiled globally; however the factors limiting a population across its migratory range are typically poorly understood. Zambia's Greater Liuwa Ecosystem (GLE) contains one of the largest remaining blue wildebeest (Connochaetes taurinus taurinus) migrations, yet the population structure, vital rates, and limiting factors are virtually unknown. We conducted a long‐term demographic study of GLE wildebeest from 2012 to 2019 of 107 collared adult females and their calves, 7352 herd observations, 12 aerial population surveys, and concurrent carnivore studies. We applied methods of vital rate estimation and survival analysis within a Bayesian estimation framework. From herd composition observations, we estimated rates of fecundity, first‐year survival, and recruitment as 68%, 56%, and 38% respectively, with pronounced interannual variation. Similar rates were estimated from calf‐detections with collared cows. Adult survival rates declined steadily from 91% at age 2 years to 61% at age 10 years thereafter dropping more sharply to 2% at age 16 years. Predation, particularly by spotted hyena, was the predominant cause of death for all wildebeest ages and focused on older animals. Starvation only accounted for 0.8% of all unbiased known natural causes of death. Mortality risk differed substantially between wet and dry season ranges, reflecting strong spatio‐temporal differences in habitat and predator densities. There was substantial evidence that mortality risk to adults was 27% higher in the wet season, and strong evidence that it was 45% higher in the migratory range where predator density was highest. The estimated vital rates were internally consistent, predicting a stable population trajectory consistent with aerial estimates. From essentially zero knowledge of GLE wildebeest dynamics, this work provides vital rates, age structure, limiting factors, and a plausible mechanism for the migratory tendency, and a robust model‐based foundation to evaluate the effects of potential restrictions in migratory range, climate change, predator–prey dynamics, and poaching.