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.

No abstract

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.

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.

IntroductionPredators can affect prey not only by killing them, but also by causing them to alter their behavior, including patterns of habitat selection. Prey can reduce the risk of predation by moving to habitats where predators are less likely to detect them, less likely to attack, or less likely to succeed. The interaction of such responses to risk with other ecological processes remains relatively unstudied, but in some cases, changes in habitat use to avoid predation may be constrained by competition: larger, dominant competitors should respond freely to predation risk, but the responses of smaller, subordinate competitors may be constrained by the responses of dominant competitors. For large grazing herbivores, an alternative hypothesis proposes that smaller prey species are vulnerable to more predators, and thus should respond more strongly to predation risk.MethodsHere, we tested these two hypotheses with 775 observations of habitat selection by four species of obligate grazers (zebra, wildebeest, puku and oribi) in the immediate presence or absence of four large carnivores (lion, spotted hyena, African wild dog and cheetah) in three ecosystems (Greater Liuwa, Greater Kafue and Luangwa Valley). Patterns of predation within this set were described by observation of 1,105 kills.ResultsOur results support the hypothesis that responses to predation risk are strongest for larger, dominant competitors. Even though zebras were killed least often, they showed the strongest shift into cover when carnivores were present. Wildebeest, puku and oribi showed weaker habitat shifts, even though they were more frequently killed. These patterns remained consistent in models that controlled for differences in the hunting mode of the predator (stalking, coursing, or intermediate) and for differences among ecosystems. There was no evidence that smaller species were subject to predation by a broader set of predators. Instead, smaller prey were killed often by smaller predators, and larger prey were killed often by larger predators.DiscussionBroadly, our results show that responses to predation risk interact with interspecific competition. Accounting for such interactions should help to explain the considerable variation in the strength of responses to predation risk that has been observed.

No abstract