Globally, large carnivores are declining due to direct persecution, habitat loss, and prey depletion. The effects of prey depletion could be amplified by changes in the composition of the herbivore (prey) community that provoke changes in carnivore diets, but this possibility has received little attention. We tested for changes over the past half‐century in prey selection by the large carnivore guild in Zambia's Kafue National Park (KNP). Across 52 predator–prey dyads, 71% of the observed changes showed that large prey have become less important and small prey have become more important. Consequently, dietary niche breadth has decreased for KNP carnivores and niche overlap has increased. We tested whether changes in the importance of prey species are related to their current abundance and uniformly found that prey that have increased in importance are now relatively common, while those that have decreased in importance are now relatively rare. We identify four potential effects of these changes for conservation (through intraguild competition, group size, the energetics of hunting, and vulnerability to snaring) that warrant investigation. Synthesis and applications. Patterns of prey selection by the large carnivores in Kafue National Park (KNP) have changed appreciably over the past half‐century. Decreased predation on large prey, which are now relatively rare, has caused niche compression and increased overlap in carnivore diets. Predation by all KNP large carnivores now concentrates on four small prey species that remain relatively abundant (impala, puku, lechwe, and warthog). Methods to detect such changes in interactions between species are well‐established, but are rarely applied to large carnivore‐ungulate systems. To guide conservation of ecosystem function, monitoring programmes should consider whether prey depletion alters the patterns of predation or competition within the predator guild because these interactions strongly affect the distribution and abundance of both predators and prey. If the patterns seen in KNP are general, then where carnivores are limited by prey depletion, conservation efforts will be most effective if they focus on mitigating the loss of large prey. In KNP, targeted efforts to protect prey larger than 200 kg, particularly buffalo, should be a priority.

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.

2020. Testing the effects of anthropogenic pressures on a diverse African herbivore community. Ecosphere 11(3):Abstract. 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 groundbased 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.

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 halfcentury 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 km 2 (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.

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