Considerations on effort, precision and accuracy for long-term monitoring of African lions (Panthera leo), when using Bayesian spatial explicit capture-recapture models, in fenced protected areas.

Intensive management is frequently required in fenced wildlife areas to reduce deleterious effects of isolation. Decisions on how best to manage such wildlife are ideally informed by regular and reliable estimates of spatiotemporal fluctuations in population size and structure. However, even in small, fenced areas, it is difficult and costly to regularly monitor key species using advanced methods. This is particularly the case for large carnivores, which typically occur at low density and are elusive yet are central to management decision-making due to their top-down effects in ecosystems and attracting tourism. In this study, we aimed to provide robust estimates of population parameters for African lions (Panthera leo) and use the data to inform a resource-efficient long-term monitoring programme. To achieve this, we used unstructured spatial sampling to collect data on lions in Pilanesberg National Park, a small (~550 km2) fenced protected area in South Africa. We used Bayesian spatial capture-recapture models to estimate density, abundance, sex ratio and home range size of lions over the age of 1 year. Finally, to provide guidance on resource requirements for regular monitoring, we rarefied our empirical data set incrementally and analysed the subsets. Lion density was estimated to be 8.8 per 100 km2 (posterior SD = 0.6), which was lower than anticipated by park management. Sex ratio was estimated close to parity (0.9♀:1♂), consistent with emerging evidence in fenced lion populations, yet discordant with unfenced populations, which are usually ~2♀:1♂ in healthy, source populations. Our rarefied data suggest that a minimum of 4000 km search effort needs to be invested in future monitoring to obtain accurate and precise estimates, while assuming similar detection rates. This study demonstrates an important utility of Bayesian spatial explicit capture-recapture methods for obtaining robust estimates of lion densities and other important parameters in fence-protected areas to inform decision-making.

Temporal variation in translocated Isle Royale wolf diet.

Wolves (Canis lupus) can exert top-down pressure and shape ecological communities through the predation of ungulates and beavers (Castor spp.). Therefore, understanding wolf foraging is critical to estimating their ecosystem-level effects. Specifically, if wolves are consumers that optimize tradeoffs between the cost and benefits of prey acquisition, changes in these factors may lead to prey-switching or negative-density dependent selection with potential consequences for community stability. For wolves, factors affecting cost and benefits include prey vulnerability, risk, reward, and availability, which can vary temporally. We described the wolf diet by the frequency of occurrence and percent biomass and characterized the diet using prey remains found in wolf scats on Isle Royale National Park, Michigan, USA, during May-October 2019 and 2020. We used logistic regression to estimate prey consumption over time. We predicted prey with temporal variation in cost (availability and/or vulnerability) such as adult moose (Alces alces), calf moose, and beaver (Castor canadensis) to vary in wolf diets. We analyzed 206 scats and identified 62% of remains as beaver, 26% as moose, and 12% as other species (birds, smaller mammals, and wolves). Adult moose were more likely to occur in wolf scats in May when moose are in poor condition following winter. The occurrence of moose calves peaked during June-mid-July following birth but before calf vulnerability declined as they matured. By contrast, beaver occurrence in wolf scat did not change over time, reflecting the importance of low-handling cost prey items for recently introduced lone or paired wolves. Our results demonstrate that the wolf diet is responsive to temporal changes in prey costs. Temporal fluctuation in diet may influence wolves' ecological role if prey respond to increased predation risk by altering foraging or breeding behavior.

The successful reintroduction of African wild dogs (Lycaon pictus) to Gorongosa National Park, Mozambique.

Large carnivores have experienced widespread extirpation and species are now threatened globally. The ecological impact of the loss of large carnivores has been prominent in Gorongosa National Park, Mozambique, after most were extirpated during the 1977-92 civil war. To remedy this, reintroductions are now being implemented in Gorongosa, initiating with endangered African wild dogs (Lycaon pictus), hereafter 'wild dogs'. We describe the first transboundary translocation and reintroduction of founding packs of wild dogs to Gorongosa over a 28-month study period and evaluate the success of the reintroduction based on five key indicator categories. We also assess how wild dog space use and diet influenced their success. We found that pre-release, artificial pack formation in holding enclosures aided group cohesion and alpha pair establishment. Post-release, we also observed natural pack formations as a result of multiple dispersal events. Founder and naturally formed packs produced pups in two of the three breeding seasons and packs successfully recruited pups. Survival rate for all wild dogs was 73% and all mortality events were from natural causes. Consequently, the population grew significantly over the study period. All indicators of success were fully achieved and this study documents the first successful reintroduction of wild dogs into a large, unfenced landscape in Mozambique and only the second on the continent. Potential mechanisms underlying these early successes were the avoidance of habitats intensively used by lions, dietary partitioning with lion, avoidance of human settlements, and Gorongosa's management strategy. We predict further population expansion in Gorongosa given that 68% of the park is still unused by wild dogs. This expansion could be stimulated by continued reintroductions over the short- to medium-term. Recovery of wild dogs in Gorongosa could aid in the re-establishment of a larger, connected population across the greater Gorongosa-Marromeu landscape.

Spatial heterogeneity facilitates carnivore coexistence.

Competitively dominant carnivore species can limit the population sizes and alter the behavior of inferior competitors. Established mechanisms that enable carnivore coexistence include spatial and temporal avoidance of dominant predator species by subordinates, and dietary niche separation. However, spatial heterogeneity across landscapes could provide inferior competitors with refuges in the form of areas with lower competitor density and/or locations that provide concealment from competitors. Here, we combine temporally overlapping telemetry data from dominant lions (Panthera leo) and subordinate African wild dogs (Lycaon pictus) with high-resolution remote sensing in an integrated step selection analysis to investigate how fine-scaled landscape heterogeneity might facilitate carnivore coexistence in South Africa's Hluhluwe-iMfolozi Park, where both predators occur at exceptionally high densities. We ask whether the primary lion-avoidance strategy of wild dogs is spatial avoidance of lions or areas frequented by lions, or if wild dogs selectively use landscape features to avoid detection by lions. Within this framework, we also test whether wild dogs rely on proactive or reactive responses to lion risk. In contrast to previous studies finding strong spatial avoidance of lions by wild dogs, we found that the primary wild dog lion-avoidance strategy was to select landscape features that aid in avoidance of lion detection. This habitat selection was routinely used by wild dogs, and especially when in areas and during times of high lion-encounter risk, suggesting a proactive response to lion risk. Our findings suggest that spatial landscape heterogeneity could represent an alternative mechanism for carnivore coexistence, especially as ever-shrinking carnivore ranges force inferior competitors into increased contact with dominant species.

Using road patrol data to identify factors associated with carnivore roadkill counts.

As the global road network expands, roads pose an emerging threat to wildlife populations. One way in which roads can affect wildlife is wildlife-vehicle collisions, which can be a significant cause of mortality through roadkill. In order to successfully mitigate these problems, it is vital to understand the factors that can explain the distribution of roadkill. Collecting the data required to enable this can be expensive and time consuming, but there is significant potential in partnering with organisations that conduct existing road patrols to obtain the necessary data. We assessed the feasibility of using roadkill data collected daily between 2014 and 2017 by road patrol staff from a private road agency on a 410 km length of the N3 road in South Africa. We modelled the relationship between a set of environmental and anthropogenic variables on the number of roadkill carcasses, using serval (Leptailurus serval) as a model species. We recorded 5.24 serval roadkill carcasses/100 km/year. The number of carcasses was related to season, the amount of wetland, and NDVI, but was not related to any of the anthropogenic variables we included. This suggests that roadkill patterns may differ greatly depending on the ecology of species of interest, but targeting mitigation measures where roads pass through wetlands may help to reduce serval roadkill. Partnering with road agencies for data collection offers powerful opportunities to identify factors related to roadkill distribution and reduce the threats posed by roads to wildlife.