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Genetic diversity is critical to a population’s ability to overcome gradual environment change. Large-bodied wildlife existing in regions with relatively high human population density are vulnerable to isolation-induced genetic drift, population bottlenecks, and loss of genetic diversity. Moose (Alces americanus americanus) in eastern North America have a complex history of drastic population changes. Current and potential threats to moose populations in this region could be exacerbated by loss of genetic diversity and connectivity among subpopulations. Existing genetic diversity, gene flow, and population clustering and fragmentation of eastern North American moose are not well quantified, while physical and anthropogenic barriers to population connectivity already exist. Here, single nucleotide polymorphism (SNP) genotyping of 507 moose spanning five northeastern U.S. states and one southeastern Canadian province indicated low diversity, with a high proportion of the genomes sharing identity-by-state, with no consistent evidence of non-random mating. Gene flow estimates indicated bidirectionality between all pairs of sampled areas, with magnitudes reflecting clustering and differentiation patterns. A Discriminant Analysis of Principal Components analysis indicated that these genotypic data were best described with four clusters and indicated connectivity across the Saint Lawrence River and Seaway, a potential physical barrier to gene flow. Tests for genetic differentiation indicated restricted gene flow between populations across the Saint Lawrence River and Seaway, and between many sampled areas facing expanding human activity. These results document current genetic variation and connectivity of moose populations in eastern North America, highlight potential challenges to current population connectivity, and identify areas for future research and conservation.

The leopard Panthera pardus is in range-wide decline, and many populations are highly threatened. Prey depletion is a major cause of global carnivore declines, but the response of leopard survival and density to this threat is unclear: by reducing the density of a dominant competitor (the lion Panthera leo) prey depletion could create both costs and benefits for subordinate competitors. We used capture–recapture models fitted to data from a 7-year camera-trap study in Kafue National Park, Zambia, to obtain baseline estimates of leopard population density and sex-specific apparent survival rates. Kafue is affected by prey depletion, and densities of large herbivores preferred by lions have declined more than the densities of smaller herbivores preferred by leopards. Lion density is consequently low. Estimates of leopard density were comparable to ecosystems with more intensive protection and favourable prey densities. However, our study site is located in an area with good ecological conditions and high levels of protection relative to other portions of the ecosystem, so extrapolating our estimates across the Park or into adjacent Game Management Areas would not be valid. Our results show that leopard density and survival within north-central Kafue remain good despite prey depletion, perhaps because (1) prey depletion has had weaker effects on preferred leopard prey compared to larger prey preferred by lions, and (2) the density of dominant competitors is consequently low. Our results show that the effects of prey depletion can be more complex than uniform decline of all large carnivore species, and warrant further investigation.

The moose (Alces alces) is a charismatic species in decline across much of their southern distribution in North America. In the northeastern United States, much of the reduction has been attributed to winter tick (Dermacentor albipictus) infestations. Winter ticks are fairly immobile throughout all life stages, and therefore their distribution patterns at any given time are shaped largely by the occurrence of moose across the landscape during the peak of two critical time periods: fall questing (when ticks latch onto moose) and spring drop-off (when engorged female ticks detach from moose). We used recent land cover and lidar data within a dynamic occupancy modeling framework to estimate first-order habitat selection (use vs. non-use) of female moose (n = 74) during the tick questing and drop-off periods. Patch extinction and colonization rates between the fall questing and spring drop-off periods were strongly influenced by habitat and elevation, but these effects were diminished during the fall questing period when moose were more active across the landscape. From the fall questing period to the spring drop-off period, patches where colonization was high and extinction was low had higher proportions of young (shrub/forage) mixed forest at higher elevations. Further, we evaluated the fitness consequences of habitat selection by adult females during the fall questing period, when females and their calves acquire ticks. We compared Resource Selection Functions (RSF) for five females that successfully reared a calf to age 1 with five females whose calves perished due to ticks. Adult female moose whose offspring perished selected habitats in the fall that spatially coincided with areas of high occupancy probability during the spring tick drop-off period. In contrast, adult female moose whose offspring survived selected areas where the probability of occupancy during the spring drop-off was low; at present, natural selection may favor female adults who do not select the same habitats in fall as in spring. Our model coefficients and mapped results define “hotspots” that are likely encouraging the deleterious effects of the tick-moose cycle. These findings fill knowledge gaps about moose habitat selection that may improve the effectiveness of management aimed at reversing declining population trends.

Lay Summary: We studied moose calves to investigate the links between stress and nutrition, a detrimental parasite during challenging winter conditions and winter survival. We identified several covariates related to fecal glucocorticoid metabolite concentrations and urea nitrogen:creatinine ratios and linked these physiological metrics to a survival rate., Moose populations in the northeastern United States have declined over the past 15 years, primarily due to the impacts of winter ticks. Research efforts have focused on the effects of winter tick infestation on moose survival and reproduction, but stress and nutritional responses to ticks and other stressors remain understudied. We examined the influence of several environmental factors on moose calf stress hormone metabolite concentrations and nutritional restriction in Vermont, USA. We collected 407 fecal and 461 snow urine samples from 84 radio-collared moose calves in the winters of 2017–2019 (January–April) to measure fecal glucocorticoid metabolites (fGCM) concentrations and urea nitrogen:creatinine (UN:C) ratios. We used generalized mixed-effects models to evaluate the influence of individual condition, winter ticks, habitat, climate and human development on stress and nutrition in calf moose. We then used these physiological data to build generalized linear models to predict calf winter survival. Calf fGCM concentrations increased with nutritional restriction and snow depth during adult winter tick engorgement. Calf UN:C ratios increased in calves with lighter weights and higher tick loads in early winter. Calf UN:C ratios also increased in individuals with home ranges composed of little deciduous forests during adult winter tick engorgement. Our predictive models estimated that winter survival was negatively related to UN:C ratios and positively related to fGCM concentrations, particularly in early winter. By late March, as winter ticks are having their greatest toll and endogenous resources become depleted, we estimated a curvilinear relationship between fGCM concentrations and survival. Our results provide novel evidence linking moose calf stress and nutrition, a problematic parasite and challenging environment and winter survival. Our findings provide a baseline to support the development of non-invasive physiological monitoring for assessing environmental impacts on moose populations.

Large carnivores are experiencing range contraction and population declines globally. Prey depletion due to illegal offtake is considered a major contributor, but the effects of prey depletion on large carnivore demography are rarely tested. We measured African lion density and tested the factors that affect survival using mark-recapture models fit to six years of data from known individuals in Kafue National Park (KNP), Zambia. KNP is affected by prey depletion, particularly for large herbivores that were preferred prey for KNP lions a half-century ago. This provides a unique opportunity to test whether variables that explain local prey density also affect lion survival. Average lion density within our study area was 3.43 individuals/100 km2 (95% CI, 2.79-4.23), which was much lower than lion density reported for another miombo ecosystem with similar vegetation structure and rainfall that was less affected by prey depletion. Despite this, comparison to other lion populations showed that age- and sex-specific survival rates for KNP lions were generally good, and factors known to correlate with local prey density had small effects on lion survival. In contrast, recruitment of cubs was poor and average pride size was small. In particular, the proportion of the population comprised of second-year cubs was low, indicating that few cubs are recruited into the subadult age class. Our findings suggest that low recruitment might be a better signal of low prey density than survival. Thus, describing a lion population's age structure in addition to average pride size may be a simple and effective method of initially evaluating whether a lion population is affected by prey depletion. These dynamics should be evaluated for other lion populations and other large carnivore species. Increased resource protection and reducing the underlying drivers of prey depletion are urgent conservation needs for lions and other large carnivores as their conservation is increasingly threatened by range contraction and population declines.

Large herbivore communities around the world have declined steeply in recent decades. Although excessive bushmeat harvesting is thought to be the primary cause of herbivore declines in many ecosystems, the direct effects of anthropogenic pressures on large herbivore populations remain poorly described in most of the systems experiencing decline. To test the extent to which large herbivores are impacted by ecological and anthropogenic factors in a protected area (PA) thought to be experiencing human‐caused decline, we fit distance sampling models to seven years of data from systematic ground‐based surveys in Kafue National Park (KNP) to estimate the population densities and distributions of 10 species of large herbivores, and to test what factors affect these parameters. Population densities of the ten most abundant large herbivores in KNP were substantially lower than those reported for an ecologically similar PA with less poaching pressure. Low densities were consistent across species and areas, though there was ecologically important variation among species and size classes. Densities of larger‐bodied herbivores were greatly depressed relative to smaller species. This pattern has direct and indirect effects on large carnivore populations, with broad implications for the ecotourism and trophy hunting industries. Statistically and methodologically rigorous methods to test the effects of anthropogenic and environmental variables on density and distribution exist, but are rarely applied to large herbivores. To quantify trends in herbivore populations and evaluate the effectiveness of conservation actions, our results show that distance sampling with stratified ground‐based monitoring is an efficient and effective method. In the Greater Kafue Ecosystem (GKE), continued increases in resource protection are needed to facilitate the recovery of an economically and ecologically important large herbivore guild. More broadly, our results confirm that anthropogenic effects on large herbivore distribution and abundance can be strong over wide areas for all species (particularly the larger members of the guild), even in very large PAs.

Protected area managers need reliable information to detect spatial and temporal trends of the species they intend to protect. This information is crucial for population monitoring, understanding ecological processes, and evaluating the effectiveness of management and conservation policies. In under-funded protected areas, managers often prioritize ungulates and carnivores for monitoring given their socio-economic value and sensitivity to human disturbance. Aircraft-based surveys are typically utilized for monitoring ungulates because they can cover large areas regardless of the terrain, but such work is expensive and subject to bias. Recently, unmanned aerial vehicles have shown great promise for ungulate monitoring, but these technologies are not yet widely available and are subject to many of the same analytical challenges associated with traditional aircraft-based surveys. Here, we explore use of inexpensive and robust distance sampling methods in Kafue National Park (KNP) (22,400 km2), carried out by government-employed game scouts. Ground-based surveys spanning 101, 5-km transects resulted in 369 ungulate group detections from 20 species. Using generalized linear models and distance sampling, we determined the environmental and anthropogenic variables influencing ungulate species richness, density, and distribution. Species richness was positively associated with permanent water and percent cover of closed woodland vegetation. Distance to permanent water had the strongest overall effect on ungulate densities, but the magnitude and direction of this effect varied by species. This ground-based approach provided a more cost-effective, unbiased, and repeatable method than aerial surveys in KNP, and could be widely implemented by local personnel across under-funded protected areas in Africa.

Conservation of competitively subordinate carnivores presents a difficult challenge because they are limited by dominant competitors. Prey depletion is one of the leading causes of large carnivore decline worldwide, but little is known about the net effect of prey depletion on subordinate carnivores when their dominant competitors are also reduced. African wild dogs are often limited by high densities of dominant competitors, particularly lions. We measured African wild dog density and survival, using mark-recapture models fit to 8 years of data from 425 known individuals in the Greater Kafue Ecosystem, Zambia. The GKE is affected by prey depletion, particularly of large herbivores, and thus the density of lions is significantly lower than ecologically comparable ecosystems. Counter to expectations from mesopredator release theory, wild dog density in GKE was far lower than comparable ecosystems with higher lion and prey density, though annual survival rates were comparable to large and stable populations. Average pack size was small and home range size was among the largest recorded. Our results show that low lion density did not competitively release the GKE wild dog population and we infer that the low density of wild dogs was a product of low prey density. Our results suggest that there is an optimal ratio of prey and competitors at which wild dogs achieve their highest densities. This finding has immediate implications for the conservation of the endangered African wild dog, and broad implications for the conservation of subordinate species affected by resource depletion and intraguild competition.

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.

Large 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.

Terrestrial large carnivores have great ecological, economic and cultural importance, but are in global decline due to habitat loss, prey depletion, poaching, retributive killing and regulated hunting. While regulated carnivore hunting potentially reduces conflict with humans and livestock, increases social tolerance and provides revenue for conservation, it can also drive population declines. Some policies regulating carnivore hunting acknowledge and address negative effects on demography and population dynamics, but others do not. Using wolves as an example, we identify four aspects of hunting policy that do not align well with ecological theory and data, and suggest resolutions for these problems that have broad relevance to exploited carnivore populations.

Estimates of population size are critical for conservation and management, but accurate estimates are difficult to obtain for many species. Noninvasive genetic methods are increasingly used to estimate population size, particularly in elusive species such as large carnivores, which are difficult to count by most other methods. In most such studies, genotypes are treated simply as unique individual identifiers. Here, we develop a new estimator of population size based on pedigree reconstruction. The estimator accounts for individuals that were directly sampled, individuals that were not sampled but whose genotype could be inferred by pedigree reconstruction, and individuals that were not detected by either of these methods. Monte Carlo simulations show that the population estimate is unbiased and precise if sampling is of sufficient intensity and duration. Simulations also identified sampling conditions that can cause the method to overestimate or underestimate true population size; we present and discuss methods to correct these potential biases. The method detected 2-21% more individuals than were directly sampled across a broad range of simulated sampling schemes. Genotypes are more than unique identifiers, and the information about relationships in a set of genotypes can improve estimates of population size.

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.

In dryland ecosystems, mobility is essential for both wildlife and people to access unpredictable and spatially heterogeneous resources, particularly in the face of climate change. Fences can prevent connectivity vital for this mobility. There are recent calls for large-scale barrier fencing interventions to address human-wildlife conflict and illegal resource extraction. Fencing has costs and benefits to people and wildlife. However, the evidence available for facilitating sound decision-making for fencing initiatives is limited, particularly for drylands. We identify six research areas that are key to informing evaluations of fencing initiatives: economics, edge permeability, reserve design, connectivity, ecosystem services and communities. Policy implications. Implementing this research agenda to evaluate fencing interventions in dryland ecosystems will enable better management and policy decisions. The United Nations Conventions on Migratory Species (CMS) and to Combat Desertification (UNCCD) are appropriate international agreements for moving this agenda forward and leading the development of policies and guidelines on fencing in drylands. Implementing this research agenda to evaluate fencing interventions in dryland ecosystems will enable better management and policy decisions. The United Nations Conventions on Migratory Species (CMS) and to Combat Desertification (UNCCD) are appropriate international agreements for moving this agenda forward and leading the development of policies and guidelines on fencing in drylands.

Packer et al. reported that fenced lion populations attain densities closer to carrying capacity than unfenced populations. However, fenced populations are often maintained above carrying capacity, and most are small. Many more lions are conserved per dollar invested in unfenced ecosystems, which avoid the ecological and economic costs of fencing.