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Most large carnivores feed on prey infrequently and may expend large amounts of energy to locate, capture and kill their prey. This makes them probabilistically vulnerable to fluctuating rates of energy acquisition over time, especially within the increasingly human-altered landscapes that dominate their remaining range. Consequently, quantifying their hunting behaviors and success rates is critical, yet direct observation of these events is rarely feasible. We theorized that we could determine prey pursuit and capture in African wild dogs(Lycaon pictus)using a mechanistic approach by constructing Boolean algorithms applied to accelerometer data derived from collar-mounted tags. Here, we used this method and then iteratively improved algorithms by testing them on observed hunts and kills of collared packs. Using this approach on 47 days of acceleration from three wild dogs in three packs, we identified 29 hunts with 10 kills, all of which were confirmed by direct observation except for a single kill. Our results demonstrate that hunting effort and success can largely be determined from acceleration data using a mechanistic approach. This is particularly valuable when such behaviors are rarely quantified and offers a template for research on foraging in canid species, while also contributing to the expanding body of literature that employs similar methods to quantify hunting in large carnivores. 

Large herbivores are in decline in much of the world, including sub-Saharan Africa, and true apex carnivores like the lion (Panthera leo) decline in parallel with their prey. As a consequence, competitively subordinate carnivores like the African wild dog (Lycaon pictus) are simultaneously experiencing a costly reduction in resources and a beneficial reduction in dominant competitors. The net effect is not intuitively obvious, but wild dogs’ density, survival, and reproduction are all low in areas that are strongly affected by prey depletion. To assess whether these correlations are causal, we tested the hypothesized mechanism, using data from 13 wild dog packs in two ecosystems to relate the energetic costs and benefits of hunting to variation in prey density, while controlling for the effects of local lion density, pack size, the number of dependent pups, and the level of protection. All of these variables affected the energetic costs and benefits of hunting. In areas with low prey density, the magnitude of movements and vectorial dynamic body acceleration (a measure of energy expenditure) both increased, the mass of killed prey decreased, and the number of kills per day did not change detectably. Programs to reduce or reverse the decline of large herbivore populations should be an effective means of improving the status of endangered subordinate competitors like the wild dog, and should be a high priority. Our results demonstrate the utility of research that integrates data from biomonitoring with direct, long-term observation of endangered species, their competitors, and their resources. 

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. 

Abstract 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 vardoniiand 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 () 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. 

Many African large carnivore populations are declining due to decline of the herbivore populations on which they depend. We recently noted that the densities of true 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 density of apex carnivores is low, so the expected effect of a simultaneous decrease in resources and dominant competitors is not obvious. We found that when prey density drops below a tipping point, the relationship of wild dog density to prey density changes sign, and wild dog density declines. We also noted that ‘prey depletion provides a mechanistically direct explanation of patterns in wild dog dynamics that have been attributed to climate change’ (Creel et al., 2023). Woodroffe et al. concur that prey depletion is an important threat, but suggest that we fail to understand the logic of their assertion that “climate change is likely to cause population collapse” (Rabaiotti et al., 2022), because the “identification of climate change as a threat is not based upon observed temporal trends in wild dog demography”. This statement misses our fundamental point. The data that Woodroffe et al. analyzed were collected over a period with rising temperatures and declining prey populations, so whether or not one tests for a time trend in demography, the data themselves are affected by two patterns: 

Within carnivore guilds, dominant competitors (e.g., lions, Panthera leo) are limited pri- marily 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 sub- stantially across sub-Saharan Africa due to human impacts, and where prey deple- tion 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 de- pletion 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 compo- nents 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 se- vere enough that its fitness costs outweigh the benefits of competitive release, such changes can fundamentally alter the balance between limiting factors for competi- tively subordinate species. 

Lions and other African large carnivores are in decline, due in part to effects of illegal hunting with snares, which can reduce prey availability and directly kill or injure carnivores. It is difficult to effectively remove snares from large ecosystems by patrolling, but an additional approach to reduce effects on large carnivores is to monitor the population closely and de-snare individuals who are found in a snare or have broken free but still carry the wire (often with serious injury). The effectiveness of de-snaring programs to reduce impacts on large carnivores has not been directly tested. Here, we used long-term demographic data from 386 individually identified lions in the Luangwa Valley Ecosystem to test the effects on population growth (λ) and population size (N) of a program to remove snares from injured lions and treat their wounds. Stochastic Leslie matrix projections for a period of five years showed that the population grew with the benefits of de-snaring, but was expected to decline without desnaring. Mean annual growth (λ) with de-snaring was 1.037 (with growth in 70% of years), closely matching observed changes in population size. Mean annual growth was 0.99 (with growth in 47% of years) for a model that assumed snared animals would have died if not treated, and 0.95 (with growth in 37% of years) for models that also accounted for super-additive effects via the death of dependent cubs and increased infanticide with increased male mortality. De-snaring requires intensive effort, but it can appreciably reduce the effect of snaring on lion population dynamics. 

Introduction: Predators 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. Methods: Here, 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. Results:Our 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. Discussion: Broadly, 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. 

Abstract 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.